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All type material of species newly described in this work has been deposited in collections of established research institutions.

Throughout this monograph, catalog numbers are cited only for type material, figured specimens or other pertinent voucher material, not for material studied in general.

To keep the volume of this publication within limits, only the collections that are holding such material are listed for each species. However, the data for the largest single architectonicid collection studied USNM , with representatives of most species discussed, have been computerized during this project, and a printed listing has been deposited in the library of the Division of Mollusks of that institution.

Other collections for which listings are available are Delaware Museum of Natural History computerized and National Museum of Wales published listing; see BIELER in TREW, The second main source of data and material came from personal field studies in South Africa and Panama, as well as from comparative studies in Bermuda in the Atlantic Ocean.

These studies, conducted in South Africa and Panama, Bermuda , concentrated on observations of the living animals, anatomy, ecology, and variability within and between populations, mainly of members of the genus Heliacus.

For gross dissections, shells were cracked and animals subsequently relaxed using magnesium chloride in distilled water or magnesium sulfate crystals "epsom salts".

Air dried shells, protoconchs, jaws and opercula were coated, and observed and photographed with a scanning electron microscope SEM, coating method and machine model depending on the electron microscope unit used.

One problem in working with Architectonicidae is that only a few species, usually from shallow water, are frequently obtained alive or are at least represented as alcohol-preserved material in collections.

Anatomical data derived from the study of this material to date serves taxonomically mainly at the generic level. Most species are known only from empty shells and comparative systematic work at the species level thus had to concentrate on shell characters.

An advantage, however, is that this largely shell-based system can be applied to fossil specimens. For the majority of the specimens studied, the following characters were observed and recorded using calipers and a dissecting microscope with a calibrated eyepiece at x magnification; mm accuracy given in parentheses : teleoconch diameter 0.

In addition, notes were compiled on characters of the periostracum and operculum, and on the shape and degree of heterostrophy of the protoconch viewed from above and, if possible, through the umbilicus; often aided by sketches made with drawing tube at 50x.

In groups with numerous similar forms e. Protoconch diameter was the largest protoconch dimension perpendicular to the columellar axis visible on the teleoconch thus reflecting slightly less than the actual larval shell diameter in tightly coiled specimens , measured from the outer comer of the varix see Fig.

Shell height was the greatest dimension parallel to the columellar axis, measured from the apex to the base of the aperture. Umbilical diameter was in ventral view the greatest distance between the columellar lip and the far side of the umbilicus, measured to the most distant tip of an umbilical crena.

The varix area, often colored dark brown, is usually recognizable even in eroded specimens. In badly eroded but important specimens, such as type material, the protoconch measurements and some characters could often be collected by viewing the protoconch through the teleoconch umbilicus.

Aberrant specimens with obviously distorted or repaired shells were measured but the results were not used in descriptions or statistics.

The terminology used for teleoconch characters, especially the various names for elements of the spiral sculpture, is based on a system originally used by BAYER fig.

For explanations see diagram, last page. General Part I. The shell shape is usually roundly cone-shaped, but occasionally coin- or disk-shaped.

The umbilicu s is a lways open, ranging from very wide to very narrow Fig. The periphery is rounded or furnished w ith one or two major keels.

The sculpture consists of more o r less finely gemmate or nodose spiral ribs. The nodules are produced by the intersection of usually weaker axial grooves with the deeper grooves between the spiral ribs.

In some forms especially Philippia and Psilaxis the spiral ribs and grooves are secondarily reduced.

Re latively smooth forms, especially members of Architectonica and Psilax is, often have a glossy shell surface and the nodules of the remaining spi ra l ribs a re usually flattened.

Only occasionally is stronger axial sculpture present. The diagram last page shows a generalized sculptural pattern in this family.

This ground plan is developed, at least initially, in all members of the family and provides excellent taxonomic characters see below. A number of major ribs and areas have been homologized throughout the family upper, lower and infra-peripheral ribs, see diagram last page and illustrations in sections on genus-group taxa , based on thei r relative position in early postlarval ontogeny and on specific qualities such as size, sculpture and coloration BIELER, a, d , a, b, , All architectonicid s show a noticeable growth mark in the initial third of the first teleoconch w ho rl, markin g the end of the early postlarval phase see Fig.

In some specimens, especially of the ge nus Heliaws, internal septa were noted Fig. X -ray photograph of the shell of Arcl1itecto11ica maxima Pu rr1, [USNM , courtesy M.

H :ir:iscwych]. Note widely open umbilicus. I N, 1. Arrow marks "stage of arrested growth"; line indicates measured protoconch size. Most archi tectonicid shells have a color pattern of more or less well- defined brown flecks at least on the peripheral ri bs.

Especially in forms from shallow water, lively color markings are present, frequently form ing regu lar patterns, flames or bands in va rious shades of brow n.

T he pigment patterns, generated by the coordinated activities of secretory cells along the length of the mantle organ, are often disturbed after repaired shell damage.

Especially in the genus Architectonica, a number of nominal species have been based on such " unique" specimens see BIELER, d, a, , and discussion under Architectonica perspectiva.

T eleoconch septation in Hclia cm implcxm M1GHELS, Specimen cut and ground to plane of penultimate whorl. It is smooth, glossy, often transparent and occasionally has a white or yellowish- to dark-brown pattern of blotches.

A varix-forming peritreme always separates the protoconch from the first teleoconch whorl. Its lower side visible on the upper side of the teleoconch has strongly bulging, inflated whorls.

Several species display a more or less distinct sculpture of axial folds in the protoconch suture Figs. A distinct sculpture of axial ribs, known from some Tertiary architectonicids BIELER, b: pl.

Some groups also have a distinct ridge on the protoconch, situated in the anal region of the larva. The ridge is referred to as the "anal keel" ROBERTSON, A few species, especially of Philippia, have a callous thickening partly or wholly overlapping the anal keel and the false umbilicus.

The functional significance of the anal keel and callus is not known. All architectonicid protoconchs are positioned at an oblique angle to the teleoconch heterostrophy, see below and are multispiral.

The measured protoconch size range of an architectonicid species usually displays a pattern of normal statistical distribution.

This phenomenon might be linked to bimodal egg size [MINNITI et al. Periostracum The periostracum consists of a relatively thin yellowish or transparent conchiolin layer, which swells strongly when wet.

In dried condition it shrinks and flakes off in whitish or brownish scales. It overlays the teleoconch sculpture, frequently enhancing sculp- tural elements such as spiral ribs and nodules by its uneven thickness.

While hiding weaker sculptural elements of the teleoconch such as axial and spiral threads, the periostracum adds its own sculpture to the overall appearance of the shell in the form of microscopic spiral sculpture.

In sand-dwelling forms, the periostracum is usually worn off and remnants of it are only found in the spiral grooves and on the umbilical wall.

In the "polyp-dwelling" Heliacus species, even fully grown specimens usually retain the periostracum, most prominently developed in H infundibuliformis see, e.

Calcareous opercula have been reported for fossil forms e. In species with relatively small, round apertures the multispiral operculum is of circular outline; tight closure is achieved by a flexible 4 5 Examples of architectonicid protoconch morphology, sketched perpendicular to teleoconch axis.

Above: as visible on shell apex; below: as visible through teleoconch umbilicus, with outline of apical aspect superimposed stippled line. GRAY in M.

GRAY, , see Fig. Architectonicid opcrcula. In larger fo rms Arcl1itecton. All archi- tectonicid opercula share a construction of spirally arranged lamellae, and a peg-like process on the bod y side, by which it is anchored to the foot muscle.

The peg can be vari ously shaped and in some g roups e. The spira l lamellae of opercula are often compressed , resulting in a flat or even overall concave shape Figs.

In some g roups e. The functional significance of this shape, w hich may be present or absent in closely related form s e. Opcrcular pegs.

Similar opercular shapes are known from several not-closely related gastropod families, such as Vermetidae Dendropoma , Hydrobiidae Gocea and Siliquariidae Tenagodus.

All of these groups also convergently share the feature of a partly uncoiled shell in some or all members; the cone-shaped operculum may be linked to that trait.

Heterostrophy The body of an architectonicid larva is dextrally organized. This is demonstrated externally by the operculum see Fig.

Despite its dextral organization, how- ever, the larval shell appears to be sinistrally coiled. The condition of having a dextrally organized body in an apparently sinistral shell, often called "hyperstrophy", occurs in several gastropod groups, e.

The protoconch apex accordingly is visible within the umbilicus of the teleoconch. Since the difference between architectonicid "anastrophy" and the "heterostrophy" of other families e.

Several reports of sinistral architectonicids can be found in the literature e. While sinistrality is a frequent phenomenon in the Gastropoda, with the regular or occasional occurrence of sinistral animals in a number of families e.

All records of "sinistral" architectonicids were found to refer to abnormal dextral hyperstrophy, whereby the teleoconch retains the hyperstrophic coiling of the protoconch.

Shifting of the mantle, resulting in a change from hyperstrophic protoconch to orthostrophic teleoconch in a "normal" architectonicid, is apparently blocked in these animals.

Other cases known are specimens of H. Larval shells unidentified Allantic architectonicid larvae. Anatomy and Biology Anatomy Previous anatomica l studies on A rchitectonicidae have bee n published by!

MERING , Bouv1rn a, b, 2 , R1sBEC , MERRILL , CuMo , and especially, H ASZPRUNAR a,b,c. ROBERTSON a, discussed the anatomical characters of th e fami ly in their relati onships to o ther groups.

The following summarizes the published informatio n, aug mented by personal observa tions mainly on species of Arcl1itectonica and Heliac11s s. The anterior portion of the foot is produced into two pointed, ve1y mobile lobes.

The sole has two gland openings, one immediately behind the anterior margin, and a much s maller one in the center regio n.

The long, tapering, very slender in living condition cephalic tentacles carry black, lens-equipped eyes at their outer bases. T he "false mouth" openi ng of the proboscis sheath opens at the tip of a short snout.

The mantle cavity comprises about half of the body whorl of the shell; the an imal is able to fully retract and tightly close the ape rture w ith its operculum.

The mantle cavity is longitudinally d ivided by a d orsal crest, built up by the posterior pedal gland , the large arterial vessel, so-called chordoid tiss ue3, long saliva ry gla nd s, and a ciliary tract at its outer edge.

This ventral ciliary tract, together w ith an opposing dorsal strip of ciliated tissue, produces a wate r current leading from left to right.

The strongly developed osphradium, with its semi-circul arly a rran ged lamellae above a large osphradial ga nglion, is situated anteriorly on the left, incurrent side.

Its morphology 2 It should be noted that the specimens studied by BoUVJrn were not "Solari trocMeare H 1NDS, ," as stated by the author, but the closely related Architectonica perspectiv a LrNNi;, The shells of the material o n which the anatomical stud ies were based Zanzibar, ROSSEAU Coll.

The foliobranch gill lamellae ROBERTSON, 4a are not homologues of the prosobranch ctenidium or the opisthobranch plicatidium MORTON, , but epithelial extensions of the well- developed hypobranchial gland.

They have no skeletal supporting rods or ciliated bands. The gill and hypobranchial gland are situated to the right of the dorsal ciliated strip, in the excurrent chamber.

The rectum and gonoducts are located on the right side of the mantle cavity, while the kidney occupies the posterior roof. The kidney is pallially situated and supplied, rather than viscerally as in "prosobranch" gastropods HAsZPRUNAR, b: The heart is positioned immediately before the proximal dorsal end of the mantle cavity.

Its atrium is anterior-left, its ventricle posterior-right in position. The pericardium communicates with the kidney lumen by a narrow ciliated duct.

The anterior right comer of the mantle cavity is occupied by the massive, glandular oviduct and the pallial vas deferens. The male has no penis.

In hermaphroditic forms see below , male, female and sperm-receptive structures are almost completely separate with independent openings.

The nervous system is distinctly streptoneurous, epiathroid, without zygoneury, and has as in the "prosobranchs" only three ganglia supra-, subintestinal-, and visceral on the long visceral loop.

The highly specialized buccal apparatus shows two main types within the family. In most groups e. The connec- tives between buccal and cerebral ganglia run within the paired proboscis protractors which in retracted condition do not pass through the cerebropedal nerve ring.

Discotectonica and Granosolarium have a short proboscis sheath, behind which the proboscis divides into a ventral part, containing a large rod-like structure see below , and a dorsal part, the esophagus proper.

Proboscis retractors are absent, their function apparently taken over by the strong longitudinal musculature of the esophagus. In all genera the esophagus is cuticularized, the stomach large and unspecialized, and the short intestine is separated from the rectum by a strong sphincter.

A dark glandular area next to the anus, close to the anterior right mantle margin, appears to represent an anal gland.

The visceral mass contains the stomach, receptaculum seminis, the unpaired digestive gland as well as testis and ovary. The columellar muscle is ventral in position and extends about three quarters of the body whorl.

The soft-body coloration results from a combination of black in preservative brownish pigment in the epidermal cells, white bodies embedded in the tissue, and to a lesser extent and only in small forms from internal organ coloration discernible through the tissue.

Species with dark shell colors usually also have darker body pigmentation. Relative to the coloration of the remaining body, the tentacles and the anterior part of the foot the body regions exposed during normal activities are most strongly pigmented; the sole and the upper head-foot areas are less strongly or not pigmented.

This and the fact that in resting position the buccal mass is considerably withdrawn, caused early workers to believe that a radula was missing in this group resulting in classifications of the Architectonicidae as an "aglossate" or "gymnoglossate" group, e.

Two main radular types are realized in the family: a five-toothed "taenioglossate-like" radula and a "ptenoglossate-like" radula with numerous marginal teeth Figs.

The first is thought to be derived from a typical taenioglossate caenogastropod radula with seven teeth per row, by loss of the pair of laterals BIELER, It is the most common radular type in the family, present in all genera but Architectonica, Adelphotectonica, Discotectonica and Granosolarium.

The "ptenoglos- sate-like" radula, present in Architectonica and Adelphotectonica, is considered second- arily derived from the five-toothed one, by multiplication of the marginals see 'Phylogeny and Fossil Record,' below.

Instead of a true radula, Discotectonica and Granosolarium have an extremely long up to one-third of the shell diameter , toothed, rod-like cuticularized structure inside a large muscular blind sac see, e.

Homology and function of this structure are unclear. The jaws are long and narrow, consisting of numerous small elements arranged in mosaic-like fashion.

The length of the jaws ranges from 0. Sex distribution Sex distribution is very variable within the family. While sexes in the Mediterranean- Atlantic species Philippia hybrida LINNE, are strictly separate MINNITI et al.

The single type of architectonicid sperm was shown to differ greatly from the "prosobranch" type. It shares several features with the Euthyneura structure of mature and developing acrosome, periodically-banded coarse fibers, modified midpiece development, pattern of nuclear condensation , while other characters separate it from that group e.

Architectonicid rndulne SEM. A probably homologous structure, a long, transversely banded column interpolated between the base of the spermatozoan nucleus and the acrosome of the mid p iece, was described for A rchitectonica perpectiva by H EA LY Nei ther of these structures was found in Psilaxis oxytropis see H EA LY, Ar chitectonicid spermatophores have been described from Heliacus.

They cons ist of lo ng mm , coiled tubes, and are handled and possibly mo lded by a sperm a- tophore groove extending o nto part of the proboscis ROBERTSON, Eggs and larval development Spawn masses are kn own fro m species of the genera Architectonica and H eliacus.

They consist of soft, gelatino us, sa usage-shaped masses, usually depend ing on animal-size several centimeters lo ng and about three millimeters in diameter.

They are ancho red to the substrate by sticky mucus. The length of the mass va ries grea tl y; ind ividuals can either prod uce o ne long contin uous mass within several hours o r severa l sho rter pieces over a period of days pers.

The weakly oval eggs abo ut 0. W ithin the a lmost transparent, viscous, mucous mass, the egg strings are covered by a closely-adhering mucous sheath.

The first cleavages occured rapidly, with embryos in the slightly older end of the egg mass considerably farther along in their development.

After three days the chalazae were largely dissolved, but the egg strings stiII interconnected by the inner mucous sheath.

After eight days the veliger stage was reached. After 18 days the first veligers hatched by actively working through the now partly-desinte- grated mucous sheath and mass.

Their size was still equivalent to the original egg size as no external food source such as nurse eggs was utilized. After 20 days the former egg mass was more or less completely dissolved, and all veligers were free.

At this stage each larva had two small velar lobes, a flat operculum, a large dark "larval organ," and a transparent, thin shell corresponding to the nucleus of the later protoconch.

Further laboratory maintenance was not successful; the larvae lived for another 20 days, without intake of the single-cell algae offered.

Planktic development of an architectonicid was described by ROBERTSON et al. The velum develops into four elongated lobes as the feeding larva grows in the plankton.

There are paired eyes but no tentacles until metamorphosis. ROBERTSON et al. According to RoBERTSON et al. After developing from between one-sixth to about one-half a whorl in one or two days, growth of the teleoconch stops.

Animals remained alive in this state of arrested growth without further changes for several months. Occasionally two or even three such marks can be seen.

Many architectonicids die at this stage: the Recent and fossil shells are common in museum collections We suggest that this high mortality is caused mainly by the spatial problems in finding hosts at this critical stage in the life cycle.

From the localities of occurrence and the maximum current velocities from the nearest potential spawning areas, ROBERTSON et al.

All members of the fam ily feed on coelenterates, and their radulae frequently ptenoglos- sate- like as in other coelenterate feeders such as E pitoniidae and alimentary system with cuticularization show several specializations.

A rchitectonica nobilis preys on actinarians. The snail rasps a hole in the base of a large actina ri an polyp, extends the proboscis into the coelenterate and continues feed ing until the prey dies BANDEL, ; see 'Habits and feeding behavior' und er A.

Members of Heliacus feed on zoanthinarians "colon ia l sea anemones," P l. C-E; ROBERTSON, ; and accounts in 'Taxonomy' section below.

The habi tat type of an architectonicid is well reflected in its shell shape; species can be grouped roughly into dwellers of sand y and those of hard substrates.

The sand-dweller, like the "sand dollar" sea urchin of similar habitats, is characterized by a depressed , shield-like shell, usually without distinct color pattern.

Large architecton- Fig. In this group belong members of the genera Discotecton- ica and Granosolarium. The other morphotype is the dweller of hard substrates such as rocks, corals and zoanthinarian polyp colonies.

It is characterized by a more or less rounded shell allowing for good maneuverability, and in shallow-water forms frequently has a camouflaging pattern.

In this group belong most species of Heliacus and Philippia. Among the predators of architectonicids are certain species of fish see, e.

At the South African shoreline of Natal, the common muricid species Morula granulata Ducws, , accounts for most of the predation on intertidal Heliacus species.

It is well adapted with its relatively small rounded shell to enter the sheltered areas in crevices and between zoanthid polyps where Heliacus lives Pl.

The muricid drills a hole in the upper part of the Heliacus shell. The reason for the internal septation found in some architectonicid shells Fig.

However, several Heliacus implexus individuals were found in Natal who had survived one or two such muricid attacks because the drill holes had led into empty chambers pers.

Recently, Luz 5 reported that the naticid Tectonatica filosa PHILIPPI, is a regular predator of the architectonicid Basisulcata lepida BAYER, in the Mediterranean.

Zoogeography The family is distributed worldwide, mainly in tropical and subtropical waters. Authors have described many local architectonicid "faunas," with different sets of nominal species, inhabiting Japan, Australia, and the African east coast.

However, as demonstrated by the maps in the 'Taxonomy' section, most Indo-Pacific species are very widely distributed, often ranging from the African east coast to the Central or even East Pacific, and many nominal species have proven to be synonyms.

For example, the "Hawaiian" species Heliacus implexus MIGHELS, is known as Heliacus codoceoae REHDER, , in the Easter Islands, as Torinista popula IREDALE, , in Australia, as Heliacus maorianus PowELL, , in New Zealand, as Heliacus homalaxis MELVILL, in India, and under its two synonyms Heliacus africanus BARTSCH, , and H.

Apparently all architectonicids have planktic veliger larvae able to drift in near-surface currents and thus to cover great distances e. The ability to delay metamorphosis allows some species even to cross the East Pacific: several species apparently extend from the Indo-West and Central Pacific to the American west coast e.

The probability of geo- graphic differentiation leading to speciation is restricted, and the wide ranges of distribution found for most architectonicids are thus explainable see, e.

The eastern Pacific fauna deserves special mention. The architectonicid species here encountered fall into three groups: 1 Indo-Pacific forms as mentioned above, 2 forms morphologically inseparable from Atlantic populations and probably isolated from them by the closure of the Isthmus of Panama in the early Pleistocene, and 3 forms that have evolved as endemic species, from either Tethys-Atlantic or Indo- Pacific stock.

Examples of amphi-American species are Architectonica nobilis and A. Some species are only known from the eastern Pacific.

However, a few species previously assumed to be endemic species to the American west coast or to have closest relationship to Atlantic forms ROBERTSON, a; KEEN, , were found to be of Indo-Pacific ancestry.

Solatisonax radialis, described from Panama, is now known from numerous localities throughout the Indo-Pacific, and eastern Pacific Heliacus bicanaliculatus is here regarded as a form of Indo- Pacific Heliacus areola, with intermediate morphs in the Marquesas and Galapagos Islands.

While most architectonicid species show little geographic variation specimens of the same species from South Africa and Hawaii, for instance, are usually indistinguisha- ble , some local forms have developed, apparently in relatively isolated areas at the fringe of the main population.

Heliacus infundibulifonnis Africa to Central Pacific and H. However, Heliacus discoideus, another locally restricted form Tuamotu Archipelago and Society Islands shows intermediate features.

Forms with characters otherwise unusual for the species are also found in the Marquesas e. The disjunct distributional patterns of some, however, do not reflect biological reality.

Especially in cases of species living in great depths, they are often merely a result of the small number of localities sampled.

Some species have only been collected by deep-water dredgings; the distribution maps will thus reflect the stations of certain expeditions working in deeper water, such as the 'ALBATROSS,' 'SIBOGA,' and 'VALDMA' cruises e.

This is used for widely geographically isolated populations believed to belong to the same species whose members are distinguished by differences in radular or teleoconch characters and for which separate names are already known in the literature e.

At present, it is unclear whether an exchange of genetic information exists between Indo-Pacific and Atlantic architectonicid populations.

This would explain the mor- phological similarity of some Atlantic and Indo-Pacific forms, which are here pro- visionally classified as subspecies.

Through climatic reasons, the only possible connec- tion allowing for larval passage is seen off the southern cape of Africa.

Following the direction of the warm Agulhas Current, the direction of larval transport here is east to west lndo-Pacific architectonicid larvae have been found in the waters southeast of the Cape of Good Hope ScHELTEMA, in litt.

Under present climatic conditions, however, there seems to be an ecological barrier: Indo-Pacific architectonicid veliger larvae can round the Cape with the Agulhas Current, but then do not find suitable habitat conditions in the South Atlantic, where the southwestern African coast is influenced by the cold Benguela Current.

The distribution of architectonicid species for which food requirements are known seems to be limited by the temperature requirements of their coelenterate prey.

In South Africa, the range of Heliacus infandibulifonnis, which prefers polyps of the zoanthid genus Jsaurus, extends into the southern Natal, the southernmost area where this zoanthid genus occurs.

Adults of the less specialized H. Architectonicid larvae have been collected at water temperatures as low as Phylogeny and Fossil Record Architectonicidae has recently attracted much attention by malacologists analyzing the systematic position of this group within the Gastropoda.

The family was classically grouped in the superfamily Cerithioidea, although MacDONALD had early pointed out the "special" position of the Architectonicidae.

Cladogram of Recent architectonicid genera modified from BIELER, fig. This was based on the peculiar combination of "prosobranch-like" and "opisthobranch-like" characters displayed by members of this group.

The similar five-toothed-taenioglossate radulae, however, might have evolved differently in these two groups, by independent reduc- tion of the outer marginal Mathildidae or lateral teeth Architectonicidae BIELER, Using the mathildid genus Gegania JEFFREYS, , as an outgroup, the family Architectonicidae was shown to be a monophyletic group that is defined by several synapomorphies in anatomical, radular, opercular, and shell characters see 'Family Architectonicidae,' below.

The most parsimonious cladogram the one that involves the fewest "ad hoc" hypotheses of homoplasy , here modified in Fig. The position of Heliacus was found to be weakly supported, the genus being mainly defined by unique derived characters and retained symplesiomorphies.

The relative branching sequence in the cladogram is supported by fossil and ontogenetic evidence BIELER, These previously recognized subfamilies Architectonicinae, Philippiinae, Heliacinae, Pseu- domalaxinae consisted of one or two nominal genera each, while other clades remained unassigned.

Subsequently, HEALY 64 stated that studies of sperma- tozoa support the subfamilial division. Architectonicid-like shells with heterostrophic protoconchs are known from as early as the Triassic Amphitomaria KoKEN, , Rinaldoconchus BANDEL, ; see BANDEL, Of the extant groups, Pseudomalaxis- and Heliacus-like shells appear earliest in the fossil record and are known from the Cretaceous, while other genera such as Granosolarium, Discotectonica, Architectonicia, and Philippia appear later, in the Eocene.

A brief overview of the sequence of appearance of architectonicid genera in time, based on fossil evidence, was given by MERRILL pl. Some groups with few extant members, such as Granosolarium and Pseudomalaxis, apparently had their major radiation in the Paleocene and Eocene.

The family is a slowly evolving group, presumably due to ongoing communication within the gene pool through teleplanic larvae see 'Zoogeography' , inhibiting geo- graphic speciation.

Comparison with the presumed sister group Mathildidae, with similar protoconch morphology, indicates that this trait was present even in their common ancestor.

Several extant species can be traced back as 'morphospecies' into the Pliocene, some e. An additional indicator for age and stability of architectonicids is the similarity of many species in the Indo-Pacific and Atlantic regions.

Genetic exchange apparently ceased in the early Pleistocene, since the existence of the Panamic isthmus, after the Atlantic and Pacific had been connected in this region for millions of years.

In the probably most conservative groups, Pseudomalaxis and Heliacus, forms can be found in either ocean that belong to the same morphospecies see Pseudomalaxis nobilis and Heliacus infundibulifonnis.

A number of pairs of very similar, corresponding species were found in the Inda-Pacific and Atlantic. A question remains concerning the driving forces behind speciation into many similar, sympatric species especially in Heliacus and Architectonica.

The investigation of South African Heliacus populations, for instance, showed no differences in microhabitat, activity period, food or reproductive biology between the common, almost equally- sized Heliacus variegatus, H areola and H trochoides.

Plate 1. Architectonica, apical aspects. Page 31 Plate 2. B: Heliacus habitat. Zoanthid zone on the South African east coast Sinkwazi, Natal, at extreme low tide.

Note several species of zoanthid "soft coral. D: Heliacus implexm MIGHELS, feeding on Palythoa natalensis CARLGREN Sinkwazi, Natal, South Africa.

Note large cone-shaped operculum. E: Heliacus trochoides DESHAYES, in feeding position on polyp of Palythoa nelliae from Reunion Rocks, N.

Note egg mass in lower left. Ar chitectonicid d eve lopment. Note chalaz ac co nnecting eggs. A DAMS, ; o ff Wai kiki, O ahu, H awaiian Island s [co urtesy D r.

Bo NAR]. B: Fu lly-developed , four-lobed veligcr stage. C : Ve liger withdrawn into shell. Note positio n of head black eye spot and velum in fro nt of it.

D: Newly metamorphosed juve nile. Note position of head eye spo t now close r to aperture. E: Newly metamorp hosed juvenile.

Note anal keel on shell a nd black " larval o rgan. F: Stage of arrested g rowth. Note colo r pattern and first part o f teleoconch whorl.

Systematic Part I. Characters used in Classification Architectonicids, and especially members of Architectonica, display very little variation in sculpture such as the number and arrangement of spiral ribs and color pattern within a species.

The spiral sculpture can be used to develop a "finger-print pattern" for each species within a group of many superficially similar forms that will identify a specimen from Africa as well as a specimen from the Central Pacific.

Teleoconch The teleoconch shape alone has no taxonomic value. Sharply keeled, coin-shaped, roundish or cone-shaped shells were convergently developed in several groups of this family.

Of value, however, is the combination of shape with certain sculptural elements. In all architectonicids certain spiral ribs and thus teleoconch areas can be homologized.

As a rule, at least in the early postlarval "arrested growth" phase, four distinct ribs are developed SSR, UMR, LMR, IPR; see diagram, last page , which differ in strength, shape, sculpture and coloration from other, secondary ribs.

A study of the position of these ribs on later teleoconch whorls show marked differences between superficially similar forms see, e.

Number and sculpture of spiral ribs were often found to be species-typical. The coloration of the teleoconch proved to be a very stable character, especially in groups with regular color patterns such as Architectonica, Philippia, and Heliacus s.

Special care needs to be taken to distinguish between normal fresh coloration and that of worn or aberrant specimens.

After shell fractures with extensive mantle tissue damage caused by wave action, fish or crab attacks , the normal sculptural and color pattern can often not be restored by the animal.

Protoconch The size range of the protoconch is species-specific, and can, in connection with other characters, be used as a taxonomic character.

It was found to be an ideal character to complement teleoconch characters, as it is not correlated with teleoconch size, not subject to pre-selection by the collector and easily measurable even in eroded specimens.

Closely related species with similar teleoconchs were frequently found to differ considerably in protoconch size.

The degree of heterostrophy, the color pattern, shape and number of whorls, presence, absence, or degree of development of umbilical folds, anal keel and callus were additional protoconch characters used in this study.

A weak edge on the protoconch appears commonly, with intraspecific variability e. Exceptions are Heliacus mighelsi and H discoideus, members of the problematic H infundibuliformis-complex.

At the species level, data are still scarce, but radular formulae have occasionally been used as additional characters to distin- guish between species-group taxa see BIELER, Data on anatomical characters, sperm, spematophores and egg masses are still too incomplete to be useful at the species level.

There is an indication that the soft-body coloration in Architectonica see Pl. A might prove to be a valuable species- specific character.

Generic Classification In this work, the established usage of subgeneric divisions in the genera Architectonica, Philippia and Pseudomalaxis has been abandoned.

I previously BIELER, demon- strated the pairs Architectonica s. Based on currently available data, the respective members of the pairs seem to be more closely related to each other than to other members in the family.

For Heliacus, the situation is different. The six subgenera here employed Heliacus s. Heliacus s. Further research may render it necessary to remove or synonymize some of its nominal subgenera, which currently are based mainly on shell characters.

Inda-Pacific Species Genus Architectonica RODING, Architectonica RODING, Type species by subsequent designation a. GRAY, , using the incorrect secondary spelling "Architectoma" : Trochus perspec- tivus LINNE, ; Recent, Inda-Pacific.

Type species by monotypy: Trochus perspectivm LINNE, Incorrect subsequent spellings: "Architectoma" J. GRAY, ; "Architeconica" STEWART, ; "Architectonia" MORCH, ; "Architec- tonium'' HAAs, ; "Soralium" SHuTo, Description Fig.

Protoconch: small to very large ca. Radula: ptenoglossate-like; rachidian if present smooth, without additional cusps, flanked by marginals on either side; outer marginals with 2 long processes each.

Operculum: horny, ear-shaped with very broad last whorl, flat with often callously thickened peg-like projection on body side. The species in this genus can be sorted into two groups, based on the sculpture of the mid-field, a part of the upper-side shell-sculpture.

The two resulting groups, here termed "perspectiva-group" and "maxima-group," are not considered to comprise monophyletic units, but such a division merely facilitates identification.

Specimens of the "perspectiva-group" have mid-rib MR areas that are undivided, while members of the "maxima-group" have mid-rib areas that are divided by a distinct groove into an upper UMR and a lower LMR mid-rib.

For a more extensive description and discussion of this genus see BIELER a: , Schematic representation of placement of major spiral ribs SSR in Architectonica, apertural aspect.

Arrow shows point of attachment of next whorl, intrageneric variation indicated by dotted lines. A; Figs. II, 7: 9, 28, pl. II, 7: 27, pl.

II, 7: 29, pl. SrnoGA Exped. Bishop Mus. Palau, 7B: , pl. Madras Gov. Seas: 10, pl. Ultramar, 59, pl. Reef Shells: 63, pl.

Shell News, J: 9, fig. Shell News, 3, fig. Ryukyu Ids. Gulf: pl. Welcome to our website The Lincoln Red is one of the oldest of the UK's Native Breeds of beef cattle and the Lincoln Red Cattle Society is a charitable organisation dedicated to the promotion of this magnificent breed.

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Lincoln Red Beef Beef Suppliers Breeder Suppliers Farm Shops Pubs and Restaurants. Furthermore, due to the pressure of competition between breeding companies, the way these techniques have been used has lead to a decline in diversity as fewer breeds and founder ancestors are used for reproduction Roughsedge et al.

Using Breeding stock from conventional agriculture directly connects organic farming with the use and consequences of these techniques.

In the legislation for organic farming it is stated that ET on the farm is forbidden Anon. So at this point organic farming is using double standards.

Moreover, organic livestock and ar- able farmers would like to establish a distinct organic breeding system, partly because of speciic requirements regarding livestock characteristics, but also because of their wish to improve the transparency of their system and products.

The organic sector still heavily depends on conventional breeding programmes Anon. Kin-breeding is a breeding method by which farmers select breeding bulls from different cow-families in their herd.

Every year these farmers select 4 to 5 bulls and use these evenly over the herd, avoiding mating of close rela- tives Baars, b; , Nauta et al.

Currently, the impact of this practice on the organic cow population in the Netherlands is minimal and organic farmers do not generally regard it as a serious alternative because kin-breeding is expected to increase inbreeding rates.

At the same time, the lack of an organic alternative seems to be an excuse for allowing the use of conventional breeding stock.

Animal breeding is not included directly in the interna- tional regulations of organic farming Anon. New legislation of organic seed production, however, is in development Baker et al.

In this paper we describe some of the possibilities and dilemmas of animal breeding in organic farming. First, an inventory was carried out to establish a clear view on the current situation of animal breeding in conventional and organic farming.

This inventory included a study of the lit- erature, interviews with ten organic farmers who were interested in livestock breeding and a ques- tionnaire distributed to all in Dutch organic dairy farmers The interviews focussed particularly on the view of farmers on using indirectly artiicial reproduction and the use of cows with a high genetic potential for production at their farms.

The questionnaire focussed on the use of AI bulls type, breed and the experience with the offspring of these bulls.

There were re- spondents, covering all types of dairy farming and years of conversion Anonymous, c.

The discussion paper also presents six different scenarios for organic breeding see Table 2. All Dutch organic dairy farmers at that time about were invited to these workshops, but only 50 farmers attended.

Each workshop started with one or two short presentations in which the main topics were ad- dressed and the structure for discussion was explained.

The discussions were held in groups of 5 to 6 persons including a discussion leader who also took notes on the discussion and the dif- ferent viewpoints.

The discussions focused on the different breeding scenarios presented in the discussion paper. Finally, the outcome of each workshop was presented and discussed in a plenary session.

At the end of a workshop the farmers were asked to ill in an additional form on the breeding scenarios they personally preferred including their arguments, and if possible to provide a time-scale or suggestions as to how their preferred breeding strategy could be implemented.

They were furthermore asked to give their preference for the organic dairy sector as a whole. The Thesis Wytze J. This paper presents the results of the choices of the farmers and the dis- cussions in the workshops.

This indicates that the farmers who attended the workshops represented a good average of the Dutch organic farms regarding farm size and cattle breeds used.

Ninety-ive per cent of all farmers liked to see a ban on the use of bulls from ET scenarios II—VI. Fifty-one per cent of all farmers preferred breeding to take place within the organic production chain scenarios IV—VI.

Overall, farmers preferred a more organic breeding strategy, more or less adapted to or based on organic farming Table 2. Farmers questioned the use of conventional breeding schemes.

They did not like the use of breeding programmes that are based on artiicial reproduction technologies and selection of high-producing animals. Secondly, also their individual motives for producing organically were an important consideration to choose for organic animal breeding.

In ET technology, hormones are used for synchronization and super-ovulation, which is in conlict with organic principles and standards.

A ban on the indirect use of ET was mentioned as a irst step in the development of a wholly organic breeding system.

Farmers did realize that a strict ban would reduce the supply of bulls used through artiicial insemination AI , especially Holstein bulls. It was unclear to them whether conventional breeding companies could produce special ET-free bulls and to what extent this would affect costs and supply.

Some of the arguments in favour of a ban on the use of ET also apply to the use of AI AI is unnatural and gametes are taken out of their natural environment.

Indeed, many livestock farmers considered the possibility of a ban on AI. However, in view of logistic and animal welfare aspects, participants recognized the absence of a practical alternative to AI.

In this case, most farmers are very pragmatic, but they also said that they have no choice, as long as there was no serious alter- native available.

Many farmers bypassed AI to some extent by keeping a young bull at the farm for serving heifers.

Among the workshop participants there were two kin-breeders of the Dutch Friesian dual- purpose breed. They actually use their own selected bulls in an on-farm kin-breeding scheme.

They got good results and consider bulls to be part of the system. The use of natural mating bulls has the additional beneits of reduced expenditure on AI and extra income from selling bulls for meat or as breeding stock.

They considered the market too small and the possible selection intensity and supply of breeding bulls too limited. They placed faith in their co-operative breeding company and in the structure of breed- ing programmes that had been used for several generations.

Many farmers did not want to give up this historic social relationship. Should breeding have to be organic, they then liked it to be organized along similar lines.

At the same time, many farmers had doubts about the value of conventional breeding for their own organic farm. Farmers had also noticed that cows from conventional breeding programmes matured too early.

They believed that this was due to selection being too strongly based on the performance of young animals, a criterion used by the breeders to shorten the generation interval.

By contrast, organic farming strives for long productive lives of animals so that the animals are used eficiently, also out of respect for integrity of animals.

Farmers therefore liked to see breeding strategies and values adapted to organic goals. Scenario II: Conventional breeding without ET Farmers continue to beneit from conventional breeding, on condition that animals used on their farm are not born from ET.

As a start for this approach, only bulls will be used which are not born from ET. Breeding organisations will be urged to work on a suficient pool of ET-free bulls of good genetic level.

Scenario III: Conventional breeding adapted to organic agriculture Breeding is based on data of performance of conventional cattle.

To overcome possible inlu- ence of G x E in estimated breeding values; 1 additive information is used for the selection of breeding stock for organic farming and 2 breeding goals are adapted to organic farming.

Scenario IV: Breeding based on organic principles The organic sector will establish its own breeding and selection of organic livestock.

The keeping and housing of the bulls is based on the rules for organic dairy farming. Scenario V: Regional breeding A selected group of breeders will provide breeding stock and semen for reproduction.

These breeders practice family or kin breeding as described by Baars b. Other farmers use bulls from these breeding farms. Scenario VI: Farm-speciic breeding Each organic farm will practice family or kin breeding.

Therefore, each farm has its own stock of breeding bulls or frozen sperm doses of these bulls. The bulls are randomly mated avoiding close relationships and inbreeding.

Cows are served naturally or inseminated artiicially. Farmers also liked to actually see the bulls so that they could personally approve of the an- imal.

Nowadays, farmers must decide on a bull on the basis of standard photos and index values. Farmers realized that this request might be dificult to honour because of international veterinary restrictions.

Smaller, local breeding companies might offer more scope in this respect. Many farmers are aware of the need to also include breeding in the organic production chain.

However, there was little consensus on how to reach this goal. Many farmers thought that the organic sector was too small for an effective breeding programme.

It was not clear how many animals would be needed for a speciically organic breeding syndicate. Another question for the development of a closed organic chain was whether the housing and management of AI bulls should be organic.

Most farmers did not have a clear view on this mat- ter. Organic housing and management for bulls are important to achieve a fully organic chain, but were not considered urgent.

Many farmers were against kin-breeding, using arguments like the danger posed by bulls, the risk of inbreeding depression, slow genetic gain and too much idealism.

It appeared that farmers in Thesis Wytze J. Some liked to learn more about it. Most farmers had become used to the large stock of selected AI bulls and could not be- lieve that selection in a small population on a farm is possible without inbreeding depression.

Farmers also complained about the distance between them and the breeding organiza- tion: they had no inluence on decision-making and there were not enough organic farmers to change this situation.

With kin-breeding and regional breeding farmers would regain control of breeding. But farmers were also afraid of putting breeding in the hands of a few kin-breeders.

So kin-breeding, if applied, should be well organized. Introducing kin-breeding on every organic dairy farm was seen as very unrealistic.

Most farmers were not breeders and kin-breeding requires special skills. In this scenario, breeding would be based only on inbreeding without crossbreeding; heterosis would not be used, which — as one farmer noted — would be rather ineficient.

Most farmers saw themselves as commercial users of genetic material rather than breeders of new outstanding genetic material. Crossbreeding between pure-bred lines from kin-breeders at com- mercial farms would create strong hybrid animals.

However, different organic farmers had different preferences, making that there was no consensus on possible breeding strategies in organic farming.

The discussion about breeding was linked to the broader discussion about developments in organic farming. In the s, organic production in the Netherlands grew rapidly as more conventional farmers converted to organic.

This resulted in an organic sector that can be seen as a relection of the conventional sector Baars, , with many highly specialized and industrial- ized features Roep, The development of the sector as a whole is aiming for more closed cycles and production chains and is therefore moving towards the agro-ecological and intrinsic approach Verhoog et al.

Figure 2. For example, a farm may have a very conventional symptom-focused approach to organic farming but may grow towards multi-functionality and dual-purpose cattle.

The challenge is to develop breeding strategies for organic farming that it organic principles in general and at the same time acknowl- edge the actual differences and dynamics of organic farming and animal breeding.

The organic sector as a whole is developing towards a more closed organic production chain, including the input of breeding material.

For example, the worldwide umbrella organization for organic farming, the International Federation of Organic Agriculture Movements IFOAM , is currently implementing the irst rules for organic plant breeding Baker et al.

Only a few years ago the topic animal breeding was included for the irst time in the programme of the IFOAM World Conference Anon.

Animal breeding for organic farming is expected to become an important issue in the near future. However, there are some important dilemmas to overcome.

Artiicial reproduction technologies - naturalness For most Dutch farmers, a ban on AI is no option. The arguments against AI are very much the same as those against ET.

Spermatozoa are taken out of their natural context, diluted and fro- zen for distribution, natural mating behaviour is excluded and more offspring per bull is created Spranger, So the choice of the Dutch organic dairy farmers is very pragmatic by considering AI as an indispensable technology.

On the other hand, some organic kin-breeders did not see natural mating and bulls on the farm as a problem. A ban on ET and AI would put the organic sector in a dificult position.

It is clear that conventional breeding companies cannot easily provide a special ET-free breeding programme for organic farming: the cost would be too high.

In the Netherlands, excluding ET would have a great impact on organic dairy farmers who use Holstein stock Anon. For these farmers the supply of AI bulls would decrease dramatically if ET by descent, i.

However, organic farming is seen as a very natural and animal friendly production system Bartussek, ; Verhoog et al. A ban on ET would safeguard organic farming from indirectly using other technologies like cloning and genetic engineering.

To overcome this dilemma, a ban on ET could be imple- Thesis Wytze J. Bapst, personal communication.

Breeding companies must consider their possibilities for developing a supply of ET-free bulls. If they cannot provide this, a special breeding programme for organic farmers would have to be developed, es- pecially for Holstein cattle.

A survey in the Netherlands yielded ET-free bulls, supplied by ive commercial breeding companies Nauta, Fifty-four per cent of these bulls still were black and white Holstein and red Holstein, breeds in which normally ET is used.

To avoid ET, farmers could also use breeds in which ET is less commonly used. However, the use of ET technologies in these breeds is on the rise.

For these breeds, the use of ET is low as they are not popular on the market. These breeds could be a starting point for ET-free breeding.

At the moment, however, there is no information available on how to cus- tomize conventional breeding values for organic farming.

In this discussion two aspects are impor- tant. The introduction of durability traits in breeding indices for conventional farming is increasing e.

Vollema, ; Van Der Beek, ; VanRaden, The second aspect is the inluence of the yet not quantiied genotype x environment interaction G x E on estimated breeding values indices for sires when considering organic and conventional farming as different environments.

Examples of special sire indices for ecological breeding are available in Germany Postler, , Wittenberg, The G x E interaction is related to the phenomenon that different genotypes of animals apparently have different levels of expression in different environments.

The magnitude of the G x E interaction can be deined as the amount of variance in P that is ex- plained by the interaction component.

In dairy cattle breeding it is common practice to ignore the G x E interaction when esti- mating breeding values of sires from data recorded within one country.

In theory this means that estimated breeding values are a composite of both Gi and Gi x Ej. In practice this is not a problem as long as recording environments are uniform and the G x E interaction explains only a small por- tion of the variance in the records observed.

However, if environments for example conventional versus organic farms interact signiicantly differently with different genotypes in the extreme situation leading to a re-ranking , either the G x E interaction is to be modelled as a correction factor when estimating breeding values from records of multiple environments or breeding values need to be estimated from and for its use in single type of environments.

The variance in perform- ance records or durability traits explained by the G x E interaction when considering conventional and organic farms as different environments is not yet quantiied Nauta et al.

In the near future, regulations for organic farming will be tightened Anon. In particular, further restrictions on the use of conventional concentrates, resulting in an increased price of organic concentrates, will result in a lower energy uptake of cattle in organic farming systems.

This will cause a further increase in the differences between conventional and organic environments and therefore will potentially increase the variance caused by the G x E interaction, especially for durability traits Buckley et al.

Given the size of the organic dairy sector Anon. At present the Dutch organic dairy sector comprises farms and about 30, lactating cows, about 20, of which are Holstein.

With such numbers, only a breeding programme for Holsteins with about 35 testing bulls — inseminations might be feasible.

If this target would be realized, it would increase the scope for organic breed- ing for other breeds too.

Today, however, we are seeing a decline in the number of farms converting to organic as markets are unstable and inancial support is limited. An international approach could solve this breeding problem.

If the sector does not grow substantially, recording information from conventional farms could be used. Recording information from only extensively managed farms could be used too, possibly minimizing disturbing effects of the G x E interaction on breeding values.

Both suggestions, especially the second one, lead to a reduction in number of records used and therefore to reduced accuracy of estimated breeding values of sires.

Another breeding strategy that is often mentioned as an opportunity for organic farming is the selection of animals with high lifetime productions Bakels, Bakels suggested that selecting for high lifetime production would automatically result in the selection of all the necessary traits for lon- gevity.

However, both, the ecological index and the longevity approach lead to longer generation intervals, and therefore slow down genetic progress as one has to wait for three or more lactations to be completed before a breeding value can be estimated.

In this way the introduction of kin-breeding on organic farms could have a positive effect on the image of organic farming. For increasing the genetic progress, the kin-breeding approach could also be used by regional breeding groups as was sug- gested in scenario V Table 2.

It seems to be worthwhile to study the possibilities of kin-breeding for organic farming on the bases of the natural character of this system.

We observed a strong wish among dairy farmers for more organic-oriented animal breeding. On the other hand, we did not ind consensus on how a more organic based animal breed- ing is to be achieved.

The development of organic breeding strategies is related to the dynamics of the organic sector, which is in the process of transition towards a production based on organic principles.

Organic breeding strategies should be in harmony with these developments. Development of organic animal breeding requires a restructuring of conventional breed- ing.

However, there are practical and institutional obstacles to overcome. At the national level, the sector is still small for an all-organic general breeding strategy with a possible exception for organic Holsteins.

For the smaller local breeds, genetic progress would be small. Here an international approach might broaden the possibilities.

Another breeding option would be a more individual approach, like with farm based kin- breeding. This would avoid the problems of using conventional breeding.

However, most farmers did not have the knowledge of kin-breeding and did not feel ready for it. The challenge seems to be how to combine the need for a farm-based selection and a collective approach.

If an international approach is chosen, the effects of the genotype x environment interaction on the estimation of breeding value may increase.

Further research is required for developing information and tools to support organic farmers in realizing organic breeding. Halberg, A systems approach to research in sustainability and organic farming.

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All rights reserved Chapter 3 Converting to organic dairy farming: consequences for production, somatic cell scores and calving interval of irst parity Holstein cows W.

Baars1, H. Bovenhuis2 1 Louis Bolk Institute, Hoofdstraat 24, NL LA Driebergen, The Netherlands 2 Animal Breeding and Genetics Group, Wageningen University, Wageningen, The Netherlands Published in Livestock Science 99 Thesis Wytze J.

Data was collected for Dutch organic farms, with a distinction made between long-standing-organic farms, converted organic farms and a reference group of conventional farms.

An animal model was used to estimate the effects of conversion on different traits based on data from converted organic farms.

Milk production was lower and somatic cell counts were higher on long-standing-organic farms than on conventional and converted organic farms.

Interestingly, at pre-organic farms, i. The estimates from our statistical analysis showed a highly signiicant decrease in milk yield and protein percentage due to conversion.

Also fat content decreased, SCS increased and AFC increased signiicantly. It can be concluded that the conversion to organic farming is a gradual process over years.

During conversion, signiicant changes in milk production, protein and fat contents and somatic cell scores took place. Age of irst calving is an important difference between organic and conventional farming.

Key words: conversion, organic farming, milk production, calving interval, somatic cell score, age at irst calving 3. In order to plan a conversion to organic production, basic information about expected changes is required.

Information that we do have is based on questionnaires Nauta et al. For our research we were able to use all the available production and fertility records from calvings from to at almost all organic dairy farms in the Netherlands, precluding the risk of selecting a speciic group of organic farms and giving us insight into the development of production and fertility over more than a decade.

Differences between organic and conventional dairy farming can be expected due to restrictions on the use of chemical fertilizer and concentrates EU, Organic roughage is produced without chemical fertilizer and consequently has lower energy and protein compounds Padel, Organic farming regulations also restrict the use of concentrates and set a limit on the content of conventional ingredients in the concentrates EU, In practice, the latter restriction will act as a inancial restraint on the use of concentrates as well.

On top of this, many organic farmers simply opt for a low input of concentrates from an organic point of view. With more organic roughage in the diet and a lower intake of energy and protein from concentrates, milk production is expected to decrease.

It is also expected that cattle with high genetic potential for production will have particular dificulty coping with organic environments Kristensen and Peder- sen, ; Nauta et al, Our interest, therefore, is directed especially at Holstein cattle.

The aim of this paper was to describe the changes prompted by conversion to organic farming, focusing on milk production traits, somatic cell score and calving interval of irst parity Holstein cows in those herds.

Data were obtained from the Dutch Herd Book and milk recording organization NRS. The data on organic farms were identiied by using the addresses of all Dutch organic dairy farms in For the organic farms also their date of conversion was known.

This information was obtained from Skal, the Dutch organic certiication organization for organic farming. In , organic dairy farms were registered with Skal.

The NRS database contained production records of of these farms. Data from conventional farming was collected by a random selection of conventional farms.

These farms were situated in the same areas as the organic farms. The data was edited in such way that it only contained records that would meet the cri- teria as described by the NRS NRS, e.

We could select 46, irst lactation records from organic farms and , irst lactation records from conventional farms. All cows calved between January and April Figure 3.

To describe the changes in breed composition per calving year of conventional, converted-to-organic and long-standing-organic farms between and we used The breed composition was calculated by adding the breed composition of each animal and dividing this by the total number of animals.

This resulted in Records for CI between irst and second calving and SCS were added to these data. The average SCS per lactation were based on somatic cell counts from test day records.

The average somatic cell count per lactation was estimated as the mean of all available test day somatic cell counts records. For SCS For CI we got The phenotypic trends were calculated as the averages per trait per calv- ing year from to This study was based on records from converted-to-organic farms only.

In addition, only daughters of sires with at least 4 offspring were selected. Models We used ASREML Gilmour et al. An animal model was used to account for genetic relationships between animals.

Pedigree information was traced back ive 2 generations and was included in the analyses. In the analyses of SCS, phenotypic observations were weighed according to the number of test day records that contributed to the mean.

The increase of HF blood at organic and conventional farms occurred mainly Thesis Wytze J. In , the long-standing- organic and converted-to-organic farms still had both about 5.

The long-standing- organic farms did have more DF blood 4. Half of the DF cattle on long-standing-organic farms were purebred DF. Between and , the contribution of other breeds on the recently converted-to-organic farms increased slightly: from 0.

Most Jersey cows were pure bred. Phenotypic trends in production, somatic cell count, calving interval and age at irst calving of irst parity Holstein cows Figure 3.

In the early nineties, when most converted-to-organic farms were still conventional, irst pa- rity Holstein cows at these farms produced about kg milk less than on the reference conventio- nal farms.

On long-standing-organic farms, irst parity Holstein cows produced approximately kg less milk than their conventional counterparts.

SE are shown in bars. Values at the y-axes are relative values. Compared to the conventional farms, long-standing-organic and converted-to-organic farms had a 0.

The phenotypic trend of al groups followed a fairly steady pattern, decreasing from 4. The percentage protein in the milk produced on long-standing-organic and converted-to- organic farms was respectively 0.

At converted-to-organic farms, protein percentage in milk declined after conversion and stabilized at the level of long-standing-organic farms.

Based on phenotypic information, long-standing-organic farms had a higher mean soma- tic cell score SCS than conventional farms.

Before conversion, the SCS on converted-to-organic farms was similar to that of conventional farms. After , the difference between converted-to- organic farms and conventional farms was 0.

Between and , the calving inter- val CI increased by about 23 days for long-standing-organic and 16 days for converted-to-organic farms. For conventional farms the increase in CI was about 12 days.

From onwards, Holstein cows on long-standing-organic farms had a higher age at irst calving AFC than on conventional or converted-to-organic farms.

Between and , when most farms converted to organic, AFC at these converted-to-organic farms increased from 26 to 27 months while AFC of conventional farms stayed at 26 months.

Effects of conversion on production, somatic cell score and fertility traits Figure 3. CI was not signiicantly inluenced by conversion. Conversion decreased milk production in irst lactations by about kg.

This decline started some years before conversion. After conversion, it took 5 years before production showed no further decline.

The biggest decline in production occurred in the period from one year before to two years after conversion, the decline being about kg.

Before conversion to organic, we found that milk fat increased. In the irst two years after conversion the percentage of milk fat decreased with 0.

After two years of conversion the percentage fat increased again. Before conversion, there had been a steady increase in protein level. From one year before to two years after conversion the protein level decreased 0.

Three years after conversion, however, protein level increased again. After con- version, SCS increased by 0. Assuming a population mean of Apparently, SCS continued to rise still after 6 years of organic production.