JOURNAL OF MORPHOLOGY 268:758–770 (2007)

The Female Genital Morphology of the Orb Weaving Agriognatha (Araneae, Tetragnathidae)

Dimitar Dimitrov,* Fernando Alvarez-Padilla, and Gustavo Hormiga

Department of Biological Sciences, The George University, Washington, D.C. 20052

ABSTRACT The female genital morphology of the spi- imply an understanding of the basic genital mor- ders in the araneoid genus Agriognatha (Tetragna- phology. Most taxonomic illustrations are done thidae) is described and illustrated. The female genitalia with the goal of depicting species diagnostic fea- of Agriognatha is characterized by a strong reduction of tures and do not necessarily provide the informa- the sperm storage organs (spermathecae) and by the pres- tion needed to understand genital morphology. ence of a specialized distal compartment of the median membranous chamber that functions as a sperm storage This is particularly true for species with complex organ (the posterior sac). The genital morphology of genital structures. This problem is also common Agriognatha species is unique among Tetragnathidae and for the descriptive work of species in which the it provides robust synapomorphic evidence for the mono- females present a low degree of sclerotization of phyly of genus. We discuss the phylogenetic implications the genitalia and do not have a well sclerotized of these new findings for the placement and monophyly epigynum (the sclerotized region that covers the of Agriognatha and for the monophyly of Tetragnathinae. internal genitalia), although there are a number of J. Morphol. 268:758–770, 2007. Ó 2007 Wiley-Liss, Inc. notable exceptions (e.g., Uhl, 1994, 2000; Michalik et al., 2005; Burger et al., 2003, 2006). It is not KEY WORDS: Tetragnathinae; spermathecae; homology; uncommon in taxonomic descriptions of the sperm storage; haplogyne; reduction females of such species to have their genital mor- phology insufficiently described and poorly illus- trated. The spider genus Agriognatha Cambridge, It has been known since the seventeenth century 1896 (Tetragnathidae) is a good example of this that different spider species differ in their genital problem. Agriognatha (Fig. 1) is a small genus of morphology. The first author to document spider Tetragnathinae known from Central and genitalia in a taxonomic work was Clerck (1757), South America. Currently ten species are known who illustrated the male palp of some, but not all, to belong in Agriognatha, but ongoing research on the species he described in his monograph on the on this genus will increase the num- Swedish spiders. Although the morphology of the ber to over 20 species (Dimitrov and Hormiga, per- female genitalia is also diagnostic for the species, sonal observation). Agriognatha females do not Clerck did not illustrate it. The morphology of cop- have an epigynum and the internal genital struc- ulatory organs in spiders is widely used now as tures are weakly sclerotized, a condition that is one of the most important diagnostic character typical of the members of the subfamily Tetragna- systems, especially at the species level (e.g., Plat- thinae. This low degree of sclerotization makes nick, 1975; Griswold, 1993; Foelix, 1996; see also morphological observations of the female genitalia Huber, 2004). The morphology of the copulatory of Agriognatha species difficult. Despite the avail- organs is also used in phylogenetic studies and in ability of female specimens in the original descrip- most of the recently published cladistic matrices it tions of several Agriognatha species the female accounts for more than half of the character data genital morphology is not discussed or illustrated (e.g., Griswold, 1990, 1993; Hormiga, 1994, 2000; at all (Bryant, 1940, 1945). Furthermore, two spe- Wang, 2002; Ramı´rez, 2003; Miller and Hormiga, 2004; Dimitrov and Ribera, in press; see Huber, 2004 for further discussion). Contract grant sponsor: U.S. National Science Foundation; Con- Despite their central role in spider systematics, tract grant numbers: DEB-0328644, EAR-0228699; Contract grant sponsors: Research Enhancement Fund and The George Washing- the copulatory organs of many species are poorly ton University. studied and their morphology is not well under- stood. This problem is not exclusive of old taxo- *Correspondence to: Dimitar Dimitrov, Department of Biological nomic treatments: it is not uncommon in modern Sciences, The George Washington University, WA, D.C. 20052. taxonomic revisions to find genital illustrations E-mail: [email protected] that are insufficient to understand the basic mor- Published online 30 May 2007 in phology of the species in question. Illustrating gen- Wiley InterScience (www.interscience.wiley.com) ital diagnostic characters does not necessarily DOI: 10.1002/jmor.10543

Ó 2007 WILEY-LISS, INC. FEMALE GENITAL MORPHOLOGY OF AGRIOGNATHA 759

Fig. 1. Agriognatha rucilla in its web. Dominican Republic. Photo by Gustavo Hormiga.

cies have been described from single female speci- type there are separate ducts for the seminal fluid mens but without providing a description or illus- to enter and leave (Austad, 1984). The haplogyne trations of the internal female genitalia (Cam- condition is primitive and consequently a symple- bridge, 1889; Mello-Leita˜o, 1947). siomorphy in Haplogynae. The entelegyne condi- Spiders in the large Araneoclada lineage (a clade tion is derived (apomorphic) and thus it provides that includes about 90% of the known spider spe- support to the monophyly of Entelegynae. Never- cies; see (Coddington et al., 2004) have been classi- theless this ‘‘division’’ is far from perfect: a number cally divided into two groups: Haplogynae and of taxa within Entelegynae have evolved haplo- Entelegynae (Simon, 1892; Wiehle, 1967; Platnick gyne genitalia (e.g., some tetragnathines; see et al., 1991; Uhl, 2002; but see Griswold et al., below) and some members of Haplogynae have fer- 2005). Support for these two groups, since they tilization ducts, such as some pholcids (Huber, were first proposed by Simon (1892), has come in 1997) or some tetrablemmids (Burger et al., 2006). part from the genital morphology. In haplogynes It seems clear that the notion that these two line- the female sperm storage organs (spermathecae) ages have somewhat uniform female genitalia is open to only one set of ducts, which function both not empirically supported (Uhl, 2002). as copulatory and fertilization ducts (or in some Most species of the family Tetragnathidae, as species, there are no ducts at all, as in the family well as the rest of araneoid spiders, have entele- ). In entelegyne spiders the spermathe- gyne genitalia (but note that the exact circum- cae open to the exterior by means of the so-called scription of Araneoidea awaits resolution; see Gris- copulatory ducts and a separate set of ducts (the wold et al., 2005). In the tetragnathid subfamily fertilization ducts) connect the spermathecae to Tetragnathinae (which includes the genera Tetrag- the uterus externus. Haplogynes and entelegynes natha, Agriognatha, , and Pachygna- have been said to have ‘‘cul-de-sac’’ and ‘‘conduit’’ tha, among others; see Hormiga et al., 1995) the spermathecae, respectively, because in the first haplogyne condition has evolved independently of type of spermathecae the seminal fluid enters and that found in the Haplogynae clade and the basal exits through the same duct and in the second araneomorph lineages: the fertilization ducts have

Journal of Morphology DOI 10.1002/jmor 760 D. DIMITROV ET AL.

Fig. 2. The internal female genitalia of Agriognatha rucilla. SEM. (A) Dorsal view. Scale bar 30 lm. (B) Close up view of the cuticle of the posterior sac. Scale bar 2 lm. (C) Lateral view. Scale bar 100 lm. (D) Copulatory duct dorso-lateral view. Scale bar 10 lm. (E) Frontal view. Scale bar 10 lm. (F) Long ductules of the copulatory duct. Scale bar 1 lm. BL booklung. been secondarily lost and there has been a general the results under certain character weighting simplification of the female genital structures. analyses suggest that they may be primitively Such simplification of the entelegyne genitalia to a haplogyne (Griswold et al., 2005: Fig. 219). Primi- secondary haplogyny condition is a rare evolution- tive haplogyny had been previously proposed for ary event. Secondary haplogyny probably evolved Archaeidae (Forster and Platnick, 1984) but in independently several times within the entele- the light of the new evidence presented by Gris- gyne group, e.g., in the families Archaeidae and wold et al. (2005) both hypotheses are plausible. Huttoniidae. However, the phylogenetic place- Despite the enormous advances brought by the ment of these two families is still unresolved and work of Griswold et al. (2005), the answer to this

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Fig. 3. The internal female genitalia of Agriognatha espanola. SEM. (A) Dorsal view. Scale bar 100 lm. (B) Close up view of the cuticle of the posterior sac. Scale bar 10 lm. (C) Close up view of the cuticle of the median sac. Scale bar 2 lm. (D) Copulatory duct, fronto-lateral view. Scale bar 10 lm. (E) Copulatory duct, close up. Scale bar 10 lm. (F) Frontal view. Scale bar. 10 lm. question requires a more resolved and robust phy- nal female genitalia exhibited a very atypical mor- logenetic hypothesis. phology. The most unusual observation, together During the course of systematic studies of tet- with the reduced spermathecae, was the presence of ragnathid spiders we have examined in some an unpaired sac that holds the sperm. The goal of detail the genital morphology of several Agriogna- this article is to describe and illustrate the peculiar tha species as well as some other closely related morphology of the female genitalia in several repre- genera. We were very surprised to discover that in sentatives of Agriognatha and to provide some com- Agriognatha species the spermathecae were so dra- ments on their possible phylogenetic significance. matically reduced that they are unlikely to be func- We did not make histological preparations and tional (as sperm storage organs) and that the inter- some questions regarding the morphology and the

Journal of Morphology DOI 10.1002/jmor 762 D. DIMITROV ET AL.

Fig. 4. The internal female genitalia of Agriognatha simoni. SEM. (A) Dorsal view. Scale bar 20 lm. (B) Close up view of the cu- ticle of the posterior sac. Scale bar 10 lm. (C) Lateral view. Scale bar 30 lm. (D) Copulatory duct, lateral view. Scale bar 10 lm. (E) Frontal view. Scale bar 10 lm. (F) Copulatory duct, close up laterally. Scale bar 10 lm.

function of the documented structures remain MATERIALS AND METHODS unanswered. Despite this, the results presented here provide a starting point for future studies on Female genitalia were dissected and the nonchitinous abdom- inal tissue was digested with SIGMA Pancreatin LP 1750 these spiders. Additionally, some of the features enzyme complex (Alvarez Padilla and Hormiga, in press), in a reported herein are documented for the first time solution of sodium borate prepared following the concentrations in araneoid spiders and we hope that they will cat- described by Dingerkus and Uhler (1977). After removing any alyze future investigations to help us better under- remaining tissues with needles, the preparations were washed stand the process of copulation and sperm transfer in distilled water and transferred to 75% ethanol. Cleaned geni- in this group. talia were observed and illustrated in 75% ethanol using a

Journal of Morphology DOI 10.1002/jmor FEMALE GENITAL MORPHOLOGY OF AGRIOGNATHA 763

Fig. 5. The internal female genitalia of Agriognatha insolita. SEM. (A) Dorsal view. Scale bar 10 lm. (B) Close up view of the cuticle of the posterior sac. Scale bar 10 lm. (C) Lateral view. Scale bar 30 lm. (D) Copulatory duct, close up. Scale bar 2 lm. (E) Tip of the copulatory duct. Scale bar 10 lm. (F) Frontal view. Scale bar 10 lm.

Leica MZ16 stereoscopic microscope with a camera lucida. High RESULTS resolution digital photos of the preparations were taken with a Nikon DXM1200F digital camera attached to a Leica MZ16A We examined specimens of four species of stereoscope. Additional details were checked with a Leica Agriognatha (A. rucilla Bryant, 1945; A. simoni DMRM compound microscope. After illustrating and photo- Bryant, 1940, A. insolita Chickering, 1956 and A. graphing the cleaned genitalia, the preparations were critically espanola Bryant, 1945). In addition we also exam- point dried and sputter coated with gold-palladium for SEM ex- ined specimens of closely related tetragnathine amination with a LEO 1430VP scanning electron microscope. taxa: versicolor (Walckenaer, 1842),

Journal of Morphology DOI 10.1002/jmor 764 D. DIMITROV ET AL.

Fig. 6. The internal female genitalia of Tetragnatha versicolor. SEM. (A) Dorsal view. Scale bar 30 lm. (B) Base of one of the spermathecae divisions. Scale bar 10 lm. (C) Frontal view. Scale bar 20 lm. (D) Lateral view. Scale bar 30 lm.

Tetragnatha sp. (from the Dominican Republic), base and are lined with numerous long-stem (McCook, 1894) and Pachygnatha gland ductules (LSD) over the cuticle (Figs. 2C, autumnalis (Marx, 1884) (see Appendix). The D,F,3D,E,F,4D,E,F,5D,E,F).Similarlyori- externally visible genital opening in all the studied ented gland ductules and cuticular structure can species is formed by a ventral lobe of the abdomi- be observed in the copulatory ducts and the base nal cuticle, which overhangs the actual opening of the spermathecae of the species in the three (which is more internal) and leads into a chamber other tetragnathine genera studied (Tetragnatha, whose dorsal wall is a continuation of the ventral Pachygnatha and Glenognatha;seeFigs.6A,B, abdominal cuticle. In all the species examined for 7D, 8B, D, 9B, D). Unlike the other genera that this study the dorsal cuticle in this area still car- we have studied, Agriognatha has spermathecae ries the abdominal setae (Se in Figs. 2A, 4C, 6D, strongly reduced in size. 9C). In Agriognatha species this chamber con- In Agriognatha species a whitish mass, consist- nects with a pair of short ducts anteriorly (the ent with a sperm mass, fills the posterior sac; copulatory ducts or ‘‘connecting ducts’’ sensu Levi, some smaller quantities of this mass were also 1980) and with another wider duct (or chamber), observed in the median sac (only in A. rucilla). which runs dorsally and divides into one or two The external cuticle of the posterior sac has sacs (two in A. rucilla, A. simoni,andA. espa- numerous short-stem gland ductules (sSD in Figs. nola;Figs.2A,C,E,4A,C,Eand3A,D,F, 2B, 3B, 4B, 5B). Such ductules are absent in the respectively) and one in A. insolita (Fig. 5A, C, uterus externus and median sacs as well as the F). These sacs are medially arranged, along the central chamber. The cuticle of the median sac is longitudinal axis of the abdomen. We have named smoother than the cuticle of the posterior one (Fig. these two sacs according to their position as me- 3C). Both sacs are of about the same size in A. dian and posterior sacs (MS and PS, respectively). rucilla.InA. simoni and A. espanola the median The copulatory ducts (CD) share a wide common sac is smaller than the posterior. As aforemen-

Journal of Morphology DOI 10.1002/jmor Fig. 7. The internal female genitalia of Tetragnatha sp. from the Dominican Republic. SEM. (A) Dorsal view. Scale bar 20 lm. (B) Posterior sac, close up. Scale bar 10 lm. (C) Frontal view. Scale bar 20 lm. (D) Spermathecae division, close up. Scale bar 10 lm.

Fig. 8. The internal female genitalia of Glenognatha foxi. SEM. (A) Dorsal view. Scale bar 20 lm. (B) Spermathecae and copulatory duct, close up dorsally. Scale bar 10 lm. (C) Lateral view. Scale bar 20 lm. (D) Copulatory duct, close up. Scale bar 1 lm. 766 D. DIMITROV ET AL.

Fig. 9. The internal female genitalia of Pachygnatha autumnalis. SEM. (A) Frontal view. Scale bar 30 lm. (B) Copulatory duct and spermathecae, lateral view. Scale bar 10 lm. (C) Dor- sal view. Scale bar 30 lm. (D) Lateral view. Scale bar 20 lm.

tioned, A. insolita does not appear to have a me- Schematic drawings of the internal female geni- dian sac. talia of Agriognatha are shown in Figure 10A, B A medially placed membranous sac without duc- for the species that do not have median sac (A. tules is found also in Pachygnatha autumnalis insolita), and Figure 10C, D for the species with (Fig. 9A, C, and D), Glenognatha foxi (Fig. 8C), median sac (A. rucilla, A. espanola and A. simoni). Tetragnatha versicolor (Fig. 6A, C, D) and Tetrag- Schematic drawings of the internal female genitalia natha sp. (Fig. 7A–C). All the studied species from in Glenognatha foxi, Pachygnatha autumnalis and these three genera have well developed spermathe- Tetragnatha versicolor are shown in Figures 11A, cae (S in Figs. 6A, B, C, 7A, C, 8A, B, 9B, D). B, 11C, D and 11D, F respectively. All studied Agriognatha species have a smaller and nearly cylindrical structure located proximally from the median sac that we have interpreted DISCUSSION as the uterus externus (UE in Figs. 2C, E, 3F, 4A, E, 5C, F). The uterus externus is smaller It is surprising that although females were than the sacs and its smooth cuticle forms a proxi- available in collections and some of them were pre- mally elongated cylindrical base, which widens viously dissected, the peculiar morphology of the at the end. Tetragnatha has a structure similar to female genitalia in Agriognatha remained unde- the posterior sac, in addition to the uterus ex- scribed. The most intriguing finding during this ternus, (Figs. 6A, C, D, 7A, B, C), while Glenog- work was the strong reduction of the spermathe- natha and Pachygnatha have only one structure cae, which is unique among tetragnathid spiders. (the uterus externus) (Figs. 8A, C and 9A, C A similar condition was reported only for Pachyg- respectively) situated between the two sperma- natha tristriata (Levi, 1980: 62, Figs. 242, 243) but thecae. this observation is dubious as the author himself

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Fig. 10. Schematic representation of the internal female genitalia in Agriogna- tha. (A) (dorsal) and (B) (lateral) represent the case in which the median sac is not developed (A. insolita). B (dorsal) and D (lateral) represent the species which have a median sac (A. rucilla, A. espanola, A. simoni). Arrow points to the posterior fold. admits: ‘‘Pachygnatha tristriata appears to lack ence of a fold of the abdomen cuticle (covered with seminal receptacles.’’ setae) that forms the female genital opening and In all other spiders spermathecae are present the somewhat flexibly attached paracymbium of except in very few species in the families Thera- the male palp could imply, not only a close phylo- phosidae, Liphistiidae, Diguetidae, Archaeidae and genetic relationship between these tetragnathine the entire family Pholcidae (Forster, 1980; Uhl, genera, but also similar copulatory mechanics. The 1994; Bertani and da Silva Junior, 2002). paracymbium in Agriognatha is not a separate The reduction of spermathecae and the presence sclerite, as opposed to the articulated condition of of a second sac lined with gland ductules and hold- other tetragnathines such as Tetragnatha, Pachyg- ing the sperm brings out questions about the natha, and Glenognatha (Hormiga et al., 1995). mechanics of copulation and sperm transfer in Nevertheless, in artificial expansions the para- Agriognatha. To answer these questions a detailed cymbium of Agriognatha moves in the same plane study of the copulation process in Agriognatha is as in the other three mentioned tetragnathine gen- needed. Nevertheless, on the basis of the knowl- era (Dimitrov and Hormiga, personal observation). edge that we have about other genera (Huber and On the basis of the latter observation we can pre- Senglet, 1997), which are thought to be closely dict that in Agriognatha the paracymbium is also related and exhibit similar morphological charac- inserted to some extent into the vulva during cop- teristics (Tetragnatha, Glenognatha, and Pachyg- ulation (Huber and Senglet, 1997). The fact that natha) we can hypothesize some aspects of copula- the lamina of the conductor in Agriognatha is tion in Agriognatha to a certain extent. The pres- closely associated with the embolus suggests that

Journal of Morphology DOI 10.1002/jmor 768 D. DIMITROV ET AL.

Fig. 11. Schematic representation of the internal female genitalia in other Tet- ragnathinae genera. Glenognatha foxi. (A) dorsal, (B) lateral. Pachygnatha autum- nales; (C) dorsal, (D) lateral. Tetragnatha versicolor; (E) dorsal, (F) lateral. Arrow points to the posterior fold. at least in some cases the conductor is inserted to- Senglet (1997), studying copulation of Tetragnatha gether with the embolus as in Tetragnatha (Huber montana, demonstrated that the conductor and Senglet, 1997). actually reached the entrance of the spermatheca. The female genitalia of all four tetragnathine Neither the conductor nor other palpal scelrites genera studied here have more or less complex were inserted in the median sac. On the other membranous unpaired structures located medially. hand, Huber and Senglet (1997) observed that the At least in Agriognatha the most distal chamber paracymbia of T. montana, T. extensa and Pachyg- (the posterior sac) appears to be responsible for natha clerki are also inserted in the genital open- sperm storage (Fig. 12A, B). The function of the ing and that it is pressed against the posterior median sac remains unknown. The presence of wall of the genital opening. membranous median sacs was previously reported An important question that directly affects the in Pachygnatha and Tetragnatha by Whiele (1963) evolutionary interpretation of the newly described and Levi (1980, 1981). Levi (1981) suggested that structures in Agriognatha is hypothesizing its ho- the ‘‘median seminal receptacles’’ are used to hold mologous counterparts in closely related taxa. The the conductor when mating. Later, Huber and small cylindrical sac situated posteriorly to the

Journal of Morphology DOI 10.1002/jmor FEMALE GENITAL MORPHOLOGY OF AGRIOGNATHA 769

Fig. 12. Digital photograph of cleared female genitalia in dorsal view of Agriognatha rucilla (A) and Agriognatha espanola (B). Scale bars 0.5 mm. uterus externus in T. versicolor and Tetragnatha terior sac in the female genitalia into a specialized sp. does not have ductules but its cuticle and posi- structure, which stores the sperm with associated tion are similar to those of the posterior sac of glands, are unique features that provide robust Agriognatha. Although in Tetragnatha the sperma- support for the monophyly of this genus. thecae are well developed and the posterior sac does not contain sperm, this sac may be the evolu- ACKNOWLEDGMENTS tionary predecessor of the posterior sac of Agriog- natha. This conjecture requires a phylogenetic Martı´n Ramı´rez provided extremely useful sugges- analysis to be tested. As a result of the reduction tions to our manuscript and Matthias Burger helped of the spermathecae and the loss of their sperm with the interpretation of some of the structures storing function, the posterior sac in Agriognatha described here. The authors also thank Suresh Benja- develops in a specialized sperm storage organ. The min, Lara Lopardo, Frederick W. Harrison and an presence of a posterior sac in Tetragnatha and anonymous reviewer for their useful comments on the Agriognatha provides additional evidence for the manuscript. Any errors or omissions are our own. We close relationship between these two genera. also thank the following curators and institutions for Recent work on the phylogenetics of Tetragna- the loan of study specimens: Norman I. Platnick, thidae places Agriognatha as a member of the sub- American Museum of Natural History (New York); family Tetragnathinae, along with the genera Gonzalo Giribet, Museum of Comparative Zoology, Pachygnatha, Tetragnatha and Glenognatha Harvard University (Cambridge); and Jonathan A. (Alvarez-Padilla, in press; see also Hormiga et al., Coddington, National Museum of Natural History, 1995). The presence of a membranous sac (or sacs) Smithsonian Institution (Washington, D.C.). in the female genitalia and the cuticular fold over the genital opening are putative synapomorphies LITERATURE CITED supporting the monophyly of tetragnathines. The insertion of the paracymbium during copulation Alvarez-Padilla F. Taxonomic revision of the spider genus Meta- might also be synapomorphic for this subfamily. bus (Araneae: Tetragnathidae) with comments on the tetrag- Although consistent with the available evidence nathid fauna of Chile and the phylogeny of Tetragnathidae. (Huber and Senglet, 1997), this latter hypothesis Zool J Linn Soc (in press). Alvarez-Padilla F, Hormiga G. A protocol for digesting internal needs to be tested with more data on the copula- soft tissues and mounting spiders for scanning electron mi- tory mechanics of additional tetragnathid species. croscopy. J Arach (in press). Although the genus Diphya appears as a member Austad SN. 1984. Evolution of sperm priority patterns in spi- of this subfamily in some of the phylogenetic anal- ders. In: Sperm competition and the evolution of mating sys- yses (Alvarez-Padilla, in press), it lacks the charac- tems, Smith RL, ed., pp. 223–249, Cambridge, Massachusetts: Harvard University Press. teristic genital morphology of tetragnathines Bertani R, da Silva Junior PI. 2002. The first mygalomorph spi- which would suggest that if Diphya is indeed a tet- der without spermathecae: Sickius longibulbi, with a revali- ragnathine, it may be a basal lineage within this dation of Sickius (Araneae, Theraphosidae, ). subfamily. J Arach 30:519–526. Bryant EB. 1940. Cuban spiders in the Museum of Comparative The apparent loss of functionality of the sperma- Zoology. Bull Mus Comp Zool Harv 86:247–554. thecae of Agriognatha, as implied by its vestigial Bryant EB. 1945. The Argiopidae of Hispaniola. Bull Mus Comp stage, coupled with the transformation of the pos- Zool Harv 95:357–422.

Journal of Morphology DOI 10.1002/jmor 770 D. DIMITROV ET AL.

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Journal of Morphology DOI 10.1002/jmor