J. Mar. Biol. Ass. U.K. (2005), 85, 291^305 Printed in the United Kingdom
Eudendrium klausi (Cnidaria, Hydrozoa), a new species of hydroid from Belize Stefania Puce*$, Carlo CerranoO, Antonio C. MarquesP and Giorgio Bavestrello* *Dipartimento di Scienze del Mare, Universita' Politecnica delle Marche, Via Brecce Bianche, I-60131 Ancona, Italy. O Dip.Te.Ris., Universita' di Genova, Corso Europa 26, I-16132 Genova, Italy. P Departamento de Zoologia, Instituto de Biocie“ncias, Universidade de Sa‹o Paulo, CP 11461, 05422-970, Sa‹o Paulo, Brazil. $ Corresponding author, e-mail:
[email protected]
A new species of the cosmopolitan genus Eudendrium, E. klausi, is described from Carrie Bow Cay, Belize. The species is characterized by a small colony with an intense blue colour, polyps armoured with large microbasic euryteles scattered on the hydranth body, hypostome, and female spadix; female gonophores on normal hydranths with a reduced number of tentacles. The new species is compared to other Eudendrium with similar cnidome, and other Caribbean species, including those described by Allman (1877). Among the latter, E. gracile, E. laxum, E. attenuatum, and E. tenellum (sensu Allman) are considered as species inquirenda due to the lack or the unsuitability of the type materials and the inconclusive original descriptions. Eudendrium exiguum, E. eximium, and E. fruticosum were also studied and demonstrated to be di¡erent from E. klausi sp. nov.
INTRODUCTION Hydroids of the Caribbean Sea and neighbouring areas have been studied by several authors, including Allman (1877), Nutting (1900, 1904, 1915), Leloup (1935), Fraser (1937, 1944, 1947), Mattox (1949), Mattox & Crowell (1951), van Gemerden-Hoogeveen (1965), Vervoort (1968), Ru«tzler (1969), Wedler (1975), Spracklin (1982), Florez Gonzalez (1983), Wedler & Larson (1986), and Calder (1991a,b). The ¢rst record of Caribbean Eudendrium was made by Allman (1877), who described seven new species from Florida Reef, Cuba, the Salt Key, and the Bahamas Bank, viz, Eudendrium attenuatum, Eudendrium exiguum, Eudendrium eximium, Eudendrium fruticosum, Eudendrium gracile, Eudendrium laxum, and Eudendrium tenellum. The degree of completeness of Allman’s (1877) descriptions is variable, but they usually lack information on nematocysts, gonosome, and sometimes even hydranths. For example, E. attenuatum and E. tenellum were described from colonies lacking hydranths and gonophores, thus making their classi¢cation in the genus Eudendrium provisional. In spite of the fragmentary descriptions, Allman’s speci¢c names were applied to di¡erent specimens collected all over the world (e.g. Ja«derholm, 1896; Ritchie, 1910; Jarvis, 1922; Hargitt, 1924, 1927; Leloup, 1937; Fraser, 1938, 1939, 1943, 1944, 1947, 1948; Yamada, 1954, 1959; Deevey, 1954; Rees & Thurs¢eld, 1965; Vervoort, 1968, 1972; Defenbaugh, 1972; Defenbaugh & Hopkins, 1973; Spracklin, 1982; Hirohito, 1988; Bouillon et al., 1995; and many more thought to have found E. tenellum, which, in fact, however is not recognizable. Of those problematic species, Spracklin (1982) listed E. attenuatum and E. eximium from Carrie Bow Cay in Belize. Journal of the Marine Biological Association of the United Kingdom (2005)
Five other species of Eudendrium, in this case adequately described, have also been reported for the Caribbean region: Eudendrium ramosum (Linnaeus, 1758) (FlorezGonzalez, 1983), Eudendrium rameum (Pallas, 1766) (Defenbaugh & Hopkins, 1973), Eudendrium capillare Alder, 1856 (Wedler, 1975; Florez-Gonzalez, 1983), Eudendrium carneum Clarke, 1882 (Vervoort, 1968; Wedler, 1975; Florez-Gonzalez, 1983), and Eudendrium bermudense Calder, 1988 (originally described for Bermuda and subsequently reported for Twin Cays, Belize; Calder, 1991a,b). In this study we describe a new species of Eudendrium, Eudendrium klausi, collected in Carrie Bow Cay, Belize, and compare it with previous species recorded for the region.
Figure 1. Eudendrium klausi sp. nov. (A) Distal part of a colony; (B) polyp; and (C) immature egg.
292
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Eudendrium klausi sp. nov. from Belize
Figure 2. Eudendrium klausi sp. nov. (A) Polyp with typical nematocyst distribution; (B) hydrocaulus with thick and uneven perisarc walls; (C) tentacle with pigment concentrated within the endodermal cells; (D) female reproductive polyp; (E) immature egg; (F) undischarged and discharged microbasic eurytele of the hydranth; and (G) arrangement of the nematocysts in the tentacle. Scales: A,B,D,E, 1 mm; C, 0.5 mm; F, 50 mm G, 60 mm. Journal of the Marine Biological Association of the United Kingdom (2005)
Eudendrium klausi sp. nov. from Belize
S. Puce et al. 293
Figure 3. Eudendrium klausi sp. nov. (A) Microphotographs of polyp; (B) hydrocaulus; (C) tentacles; (D) immature eggs; and (E) nematocysts in a tentacle. Journal of the Marine Biological Association of the United Kingdom (2005)
294
E. angustum
E. arbuscula
E. bathyalis
E. caraiuru
E. eximium
E. fruticosum
E. generale
Author
sp. nov.
Warren, 1908
Wright, 1859
Allman, 1877
Fascicled: no
Fascicled: no
Fascicled: yes
Marques & Oliveira, 2003 Fascicled: no
Allman, 1877
Colony
Marques & Calder, 2000 Fascicled: slightly from the base to half of the colony
Fascicled: strongly in the basal part of the colony
Fascicled: strongly up to half of the colony
Maximum height: 30 mm Branches: irregular, radial
Maximum height: 76 mm Branches: irregular
Maximum height: 45 mm Branches: irregular, radial
Maximum height: 18 mm Branches: rare, irregular, radial
Maximum height: 54 mm Branches: irregular, radial
Maximum height: 50 mm Branches: irregular, radial
Remarks: gnarled hydrocaulus of stem
Remarks: none
Remarks: none
Remarks: none
Number: 3^8
Number: 2^10
Zone: base of several branches
Zone: base of branches and irregularly elsewhere
Number: throughout Zone: throughout
Remarks: branches on all the stem up to third order, fascicled region up to 0.18 mm in diameter Number: 2^5
Maximum height: 150 mm Branches: somewhat alternate, almost planar Remarks: fascicled region up to 3 mm in diameter
von Lendenfeld, 1885 Fascicled: yes, varying from lightly to strongly, depending on the size of the colony Maximum height: 40 mm Branches: irregular, radial
Remarks: none
Remarks: none
Annulations
Stem: 0.20^0.25 Stem: 41 mm Branches: 0.13^0.22 Branches: 0.38 Pedicel: 0.15 Pedicel: 0.19 Hydranth measure- Maximum height: Maximum height: ments (mm) 0.70 0.31 Diameter: Diameter: 0.5^0.6 0.29 Remarks: none Remarks: bearing blue pigment granules in the gastrodermis
Trophosome diameters (mm)
Remarks: none
Zone: unfascicled portions, main stem and base of branches and pedicels Remarks: none
Stem: n.d. Stem: 0.08^0.10 Branches: n.d. Branches: 0.06^0.08 Pedicel: n.d. Pedicel: 0.05^0.06 Maximum height: Maximum height: 0.5^0.6 0.1^0.25 Diameter: Diameter: n.d. 0.1^0.29 Remarks: none Remarks: band of large nematocysts just above a basal annular incision
Number: 3^8
Number: 2^5 (rare)
Remarks: fascicled region up to 0.7 mm in diameter
Remarks: none
Number: 2^10 Number: 3^20 (rare) Zone: unfascicled Zone: unfascicled Zone: unfascicled Zone: unfascicled portions, main stem portions, main stem portions, main stem portions, main stem and base of and base of and base of and base of branches and branches and branches and branches and pedicels pedicels pedicels pedicels Remarks: none Remarks: none Remarks: none Remarks: distal parts of unfascicled stems indistinctly corrugated Stem: 0.40 Stem: 0.42^0.48 Stem: 0.21^0.23 Stem: 0.12^0.15 Branches: 0.20^0.25 Branches: 0.26^0.28 Branches: 0.12^0.23 Branches: 0.12^0.15 Pedicel: 0.10 Pedicels: 0.15^0.25 Pedicel: 0.12^0.15 Pedicel: 0.10^0.15 Maximum height: Maximum height: Maximum height: Maximum height: 0.18^0.75 0.55 0.50 n.d. Diameter: Diameter: Diameter: Diameter: 0.18^0.57 0.40 0.33^0.40 0.20^0.30 Remarks: poorly Remarks: none Remarks: none Remarks: preserved nematocyst pads on hydranth body, sometimes forming a continuous belt
Eudendrium klausi sp. nov. from Belize
E. klausi
S. Puce et al.
Journal of the Marine Biological Association of the United Kingdom (2005)
Table 1. Synopsis of characteristics of Eudendrium species with similar cnidome as E. klausi sp. nov.; n.d., no data.
Journal of the Marine Biological Association of the United Kingdom (2005)
E. klausi Hypostome measurements (mm)
Maximum height: 0.1^0.2 Diameter: 0.3^0.4 Remarks: bearing blue pigment granules in the gastrodermis
Tentacles
Number: 27^30 Remarks: spiny and bearing blue pigment granules regularly distributed in the endodermal cells Clusters: 6^7
Female gonophores
E. arbuscula
Maximum height: Maximum height: 0.16 n.d. Diameter: n.d. Diameter: n.d. Remarks: none Remarks: blocked by a plug of elongated endoderm cells continuous with the digestive endoderm at the base of hydranth Number: 25^30 Number: 20 Remarks: none Remarks: none
Unknown
Clusters: n.d.
E. bathyalis
E. caraiuru
E. eximium
E. fruticosum
E. generale
n.d.
n.d.
n.d.
Maximum height: 0.25 Diameter: 0.26 Remarks: none
n.d.
Number: 18^24 Remarks: none
Number: 23^34 Remarks: none
Number: 20 Remarks: none
Number: 20^27 Remarks: none
Number: 18^30 Remarks: none
Clusters: 3^5
Clusters: 4^7
Clusters: not Clusters: 6^12 Clusters: 4^5 clearly visible (15 when mature) Blastostyle: normal Blastostyle: normal Blastostyle: normal Blastostyle: Blastostyle: Blastostyle: or slightly reduced completely reduced completely reduced completely reduced in the tentacles during ontogeny during ontogeny during ontogeny
Spadix: absent Spadix: immature unbranched, mature shed Remarks: mature Remarks: mature eggs almost eggs oval, located at some distance on circular enveloped by a gelatinous the pedicel ¢lm, and linked to the hydranth body by long peduncles
Spadix: immature unbranched, mature shed Remarks: gonophores orange
(Continued)
S. Puce et al. 295
Spadix: immature Spadix: immature unbranched, unbranched, mature shed mature shed Remarks: later Remarks: vaseRemarks: mature shaped pedicels and stages of blastostyle eggs oval, continue to grow eggs completely encapsulated by ahead of the enveloped by a a perisarc layer, gelatinous capsule, gonophores, or it scattered on the may be shed scattered on the pedicel or in a pedicel or in a circle around circle around hydranth body, hydranth body, linked by short linked by short peduncles to basal peduncles to basal part of hydranth part of hydranth body or direct to body or direct to pedicel pedicel; mature eggs oval Spadix: immature unbranched
Eudendrium klausi sp. nov. from Belize
Blastostyle: immature with unreduced hypostome but with 20^22 tentacles Spadix: immature unbranched Remarks: completely wrinkled vase-shaped pedicel, spadix bearing blue pigment granules in the gastrodermis
E. angustum
296
E. angustum
E. arbuscula
E. bathyalis
E. caraiuru
E. eximium
E. fruticosum
Male gonophores
Unknown
Unknown
Clusters: n.d. Chambers: 2 Blastostyles: completely reduced
Unknown
Clusters: 10^29 Chambers: 1^2 Blastostyles: completely reduced
Unknown
Clusters: 7^14 Chambers: 2^3 Blastostyles: normal
Large microbasic euryteles
Shape: bean shaped
Shape: n.d.
Size: 23.312 mm
Size 23.310.4 mm
Size: n.d.
Size: 22.1^23.4 8.2^8.9 mm
Size: 18.7^22.7 7.1^9.3 mm
Size: 15.0^19.1 7.5^9.1 mm
Size: 18^207.5^8 mm
Distribution: very abundant at the base and scattered on other portions of the hydranth, rare on hypostome, abundant on female spadix, and coenosarc Remarks: none
Distribution: hypostome and body of hydranth
Distribution: hydranth body, male sporosac
Distribution: common on hydranth body, rare on egg peduncle, coenosarc
Distribution: common on hydranth body, hypostome, coenosarc, terminal tubercle, and immature female spadix
Distribution: rare at the base of hydranth, coenosarc, and (?)hypostome
Remarks: none
Remarks: none
Remarks: none
Remarks: none
Remarks: none
Belize
South Africa, East Madagascar Warren, 1908; Millard, 1975
North Seas
Bermuda
Brazil
Florida (USA)
Calder, 1972; Marques et al., 2000a
Marques & Calder, 2000
Marques & Oliveira, 2003
Allman, 1877; this study
Distribution Source
This study
E. generale
Clusters: up to 20 Chambers: 2^3 Blastostyles: tentacles completely absent in all stages Remarks: Remarks: Remarks: distal Remarks: apical immature distal gonophores orange chamber with pad of nematocysts terminal tubercle chamber often with on terminal a knob lost at but lacking apical tubercle maturity nematocyst pad Shape: n.d. Shape: bean shaped Shape: bean shaped Shape: bean shaped Shape: bean shaped Shape: bean shaped Size: 12^157^9 mm; also 21^2310^12 mm Distribution: Distribution: abunmoderately dant at the base of common to rare on hydranth body, rare on hypostome, hydranth body and nematocysts ring rare on spadix of immature female when present, also gonophores, and on small clusters on female spadix coenosarc Remarks: none
Remarks: shaft with few spines and discharged sideways Florida (USA) Australia, Antarctica Allman, 1877; this Watson, 1985; Puce study et al., 2002
Eudendrium klausi sp. nov. from Belize
E. klausi
S. Puce et al.
Journal of the Marine Biological Association of the United Kingdom (2005)
Table 1. (Continued.)
Journal of the Marine Biological Association of the United Kingdom (2005)
E. minutum
E. moulouyensis
E. novaezelandiae
E. pocaruquarum
E. rameum
E. ramosum
Author
Watson, 1985
Watson, 1985
Watson, 1985
Marques, 1995
(Pallas, 1766)
(Linnaeus, 1758)
Colony
Fascicled: no
Fascicled: no
Fascicled: no
Marques, Pe•a Cantero, Vervoort, 2000 Fascicled: no
Fascicled: no
Fascicled: very
Fascicled: no
Maximum height: 56 mm Branches: irregular, radial
Maximum height: 120 mm Branches: irregular, radial
Maximum height: 44 mm Branches: irregular, radial
Remarks: none
Remarks: none
Remarks: none
Number: throughout Zone: throughout
Number: n.d.
Number: 3^12
Stem: 0.15^0.18 Branches: n.d. Pedicels: n.d.
Stem: 0.26 Branches: n.d. Pedicels: n.d.
Maximum height: 0.57 Diameter: 0.18^0.39
n.d.
Maximum height: Maximum height: 43 mm 28 mm Branches: scatBranches: tered, irregular, irregularly radial; rarely alteralternate, more nate or less in one plane, upward directed Remarks: none Remarks: none
Annulation
Number: n.d.
Number: 3^12
Zone: unfascicled Zone: unfascicled portions, main stem portions, main stem and base of and base of branches and branches pedicels
Maximum height: 3 mm Branches: simple or few branched, irregularly
Maximum height: 15 mm Branches: irregular, radial
MarktannerTurneretscher, 1890 Fascicled: yes, almost to the tip of the colony Maximum height: 60 mm Branches: irregular, radial
Remarks: pedicels arise direct from stem Number: 10 (branches) Zone: proximal part of the branches, pedicels annulated throughout
Remarks: pedicels arise direct from stem Number: 3^4
Remarks: distal branches roughly alternate Number: 7
Zone: at origin of pedicels
Zone: unfascicled Zone: unfascicled Zone: unfascicled portions, main stem portions, main stem portions, main stem and base of and base of and base of branches and branches, pedicels branches, pedicels pedicels annulated annulated throughout throughout Remarks: none Remarks: none Remarks: none Remarks: stems are rearely not annulated throughout
Remarks: none
Remarks: none
Remarks: spiral annulation
Stem: 0.18^0.20 Branches: n.d. Pedicel: 0.10^0.13 basally, 0.13^0.18 apically
Stem: 0.08^0.11 Branches: 0.08^0.1 Pedicel 0.08^0.10
Stem: 0.075 Branches: n.d. Pedicel: 0.075^0.1
Remarks: sometimes corrugated at the base of pedicels Stem: 0.12^0.15 Stem: 0.7 Branches: n.d. Branches: 0.1^0.13 Pedicels: 0.11^0.13 Pedicels: 0.13^0.15
Hydranth measure- Maximum height: ment (mm) n.d. Diameter: n.d.
Maximum height: 0.70 Diameter: 0.13^0.60
Maximum height: n.d. Diameter: 0.24^0.25
Maximum height: 0.68 Diameter: 0.32
Trophosome diameter (mm)
Maximum height: n.d. Diameter: 0.13^0.15
Stem: 0.20^0.25 Branches: 0.20^0.25 Pedicel: 0.13^0.15 basally, 0.20^0.25 apically (not all pedicels vase-shaped) Maximum height: 0.90 Diameter: 0.27^0.60
(Continued)
S. Puce et al. 297
E. merulum
Eudendrium klausi sp. nov. from Belize
E. kirkpatricki
298
E. merulum
E. minutum
E. moulouyensis
E. novaezelandiae
E. pocaruquarum
E. rameum
E. ramosum
Hydranth measurement (mm)
Remarks: none
Remarks: none
Remarks: none
Remarks: none
Remarks: none
n.d.
Remarks: none
Hypostome measurements (mm)
n.d.
n.d.
n.d.
Remarks: proximal groove in the aboral region absent; zooxanthellae in the hydranth and in the coenosarc n.d.
n.d.
n.d.
n.d.
n.d.
Number: 20
Number: 25^28
Number: 20^24
Number: 17^27
Number: 20
Number: 22^35
Tentacles
Number: poorly Number: 24 preserved Remarks: none Remarks: none Female gonophores Clusters: 3^6 Clusters: up to 8 Blastostyle: Blastostyle: completely reduced completely reduced during ontogeny during ontogeny Spadix: immature unbranched, mature shed Remarks: mature eggs oval, remaining in the original position
Spadix: immature unbranched, mature shed Remarks: immature gonopores diskshaped, mature eggs scattered along a wrinkled blastostyle
Remarks: none Unknown
Remarks: none Remarks: none Remarks: none Remarks: none Remarks: none Clusters: 2 Clusters: up to 5 Clusters: 3^5 Clusters: n.d. Clusters: 3^15 Blastostyle: normal Blastostyle: reducBlastostyle: Blastostyle: Blastostyle: tion of tentacle size completely reduced completely reduced completely reduced and sometimes during ontogeny during ontogeny during ontogeny number Spadix: immature Spadix: immature Spadix: immature Spadix: immature Spadix: immature unbranched, unbranched, unbranched, unbranched, unknown, mature mature shed mature shed mature shed mature shed shed Remarks: mature Remarks: mature Remarks: mature Remarks: single Remarks: mature eggs encapsuled by egg enclosed in a encapsuled by thin eggs distributed on eggs encapsuled by a gelatinous/ the pedicel gelatinous layer, thick transparent a perisarc layer, perisarcal layer, distributed in a pellicle sessile, distributed sessile, distributed whorl around over blastostyle over blastostyle or blastostyle at their original position around blastostyle
Eudendrium klausi sp. nov. from Belize
E. kirkpatricki
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Table 1. (Continued.)
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Male gonophores
E. kirkpatricki
E. merulum
E. minutum
E. moulouyensis
E. novaezelandiae
E. pocaruquarum
E. rameum
E. ramosum
Clusters: n.d. Chambers: 2 Blastostyles: normal
Clusters: up to 28 Chambers: 2^3 Blastostyles: completely reduced
Unknown
Unknown
Clusters: up to 12 Chambers: 1^2 Blastostyles: completely reduced
Clusters: 4^8 Chambers: 2 Blastostyles: normal
Clusters: n.d. Chambers: 1^2 Blastostyles: normal
Remarks: none
Remarks: none
Remarks: none
Clusters: 3^6 Chambers: 3^4 Blastostyles: reduction of tentacle size or fully developed blastostyles Remarks: terminal tubercle lost at maturity
Remarks: none
Large microbasic euryteles
Australia, Papua New Guinea
Australia, Mediterranean Sea, Brazil
Australia
Chafarinas Islands (Mediterranean Sea)
New Zealand
Brazil
Watson, 1985
Watson, 1985; Bavestrello & Piraino, 1991; Marques, 2001; Marques et al., 2000b
Watson, 1985
Marques et al., 2000b
Watson, 1987; Schuchert, 1996
Marques, 1995, 2001
Mediterranean Sea, Atlantic Ocean, Indian Ocean Millard, 1977; Marques et al., 2000b
Shape: bean shaped
Size: 22.2^43 8.7^18 mm Distribution: rare on hydranth body, abundant on hypostome, and female spadix Remarks: shaft discharging sideways, distal end in£ated and spirally annulated, with numerous bristles arising from the annulations, thread very long with distinct spines Cosmopolitan
Watson, 1985; Marques et al., 2000b; Marques, 2001
S. Puce et al. 299
Source
Shape: (1) bean Shape: bean shaped Shape: bean shaped Shape: bean shaped Shape: bean shaped shaped; (2) elongated, paddleshaped Size: 12.8^19.2 Size: 17^25.8 Size: 10^15 Size: (1) 17^31 Size: 29^30 Size: 16.5^25 Size: (1) 21^22 5.4^8.0 mm 8.9^10.8 mm 5.0^8.7 mm 8^15 mm; 13^15 mm 8^13.3 mm 10^13mm; (2) 11^17 (2) 18^197 mm 4.5^8.5 mm Distribution: n.d. Distribution: Distribution: Distribution: Distribution: Distribution: Distribution: hydranth body, (1) abundant on sparingly over the (1) moderately hydranth body, moderately abunhypostome, and nematocyst ring, hydranth body, common on dant around hypos- terminal tubercle of female spadix few on hypostome, hypostome and hypostome; male sporosac, and tome gonophores, and coenosarc (2) abundant female spadix coenosarc; (2) around hypostome female gonophores Remarks: none Remarks: none Remarks: (1) Remarks: shaft Remarks: none Remarks: shaft Remarks: (1) discharged at 45‡ thick, crossing ca. very ornamented discharges sideangle to main axis, two-thirds the ways, thread thick, thread with ¢ne length of with spirals of ¢ne bristles undischarged bristles; (2) capsule numerous spines on distal neck region and ahead, thread ¢ne
Eudendrium klausi sp. nov. from Belize
Distribution
Remarks: wrinckled pedicel, chambers connected by a distinct neck Shape: bean shaped Shape: bean shaped
300
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Eudendrium klausi sp. nov. from Belize
Figure 4. Eudendrium fruticosum Allman, 1877. (A) Female reproductive polyp; and (B) female gonophore enveloped by a gelatinous capsule. Scales: A, 0.8 mm; B, 0.2 mm.
MATERIALS AND METHODS
RESULTS
The specimen of Eudendrium klausi was collected on apical branches devoid of coenenchyme of the sea fan Pseudopterogorgia bipinnata (Verrill, 1864), at a depth of 15 m on the south side of Carrie Bow Cay reef, Belize (see Ru«tzler & Macintyre, 1982 for a description of the area) during a research expedition carried out in July 2001. The collected sample were ¢xed in 4% formaldehyde. Drawings and microphotographs of the ¢xed material were made under low and high power microscopes. The cnidome terminology follows Weill (1934) and Mariscal (1974), and measurements of each type of nematocyst were made on ten non-discharged capsules of the ¢xed material. The specimen herein described was compared to Allman’s type specimens borrowed from the Museum of Comparative Zoology of Harvard University (MCZ) and to literature data. Type material of the new species was deposited in the United States National Museum (Natural History).
SYSTEMATICS Family EUDENDRIIDAE L. Agassiz, 1862 Genus Eudendrium Ehrenberg, 1834 Eudendrium klausi sp. nov. (Figures 1^3)
Journal of the Marine Biological Association of the United Kingdom (2005)
Type material
Holotype: female colony consisting of erect stems growing on Pseudopterogorgia bipinnata (Verrill, 1864). (South side of Carrie Bow Cay reef (Belize); water depth: 15 m). [USNM 1025073]. Collected by C. Cerrano, July 2001. Diagnosis
Eudendrium with unfascicled stems, hypostome large and £ared, tentacles 27^30 in number. Nematocyst distribution on the tentacles giving them a spiny aspect. Female blastostyles styloids, with one simple and curved spadix
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S. Puce et al. 301
Figure 5. Eudendrium fruticosum Allman, 1877. (A) Microphotographs of polyp; (B) male reproductive polyp; (C) female reproductive polyp; (D) egg with spadix enveloped by a gelatinous capsule; (E) perisarc cup persisting after the planula release; and (F) nematocysts in the stem. Journal of the Marine Biological Association of the United Kingdom (2005)
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Eudendrium klausi sp. nov. from Belize
per egg, tentacle number of blastostyles reduced. Hydranth body, hypostome, tentacles, coenosarc, and spadix deep blue. Small microbasic euryteles distributed over hydranth body, hypostome, coenosarc, regularly on tentacles. Large microbasic euryteles distributed over hydranth body, hypostome, coenosarc and female spadix. Description
Colony dioecious, up to 30 mm in height; main stems monosiphonic. Numerous stems arising from creeping hydrorhiza growing over gorgonian skeleton (Figure 1A); branches few, irregular, occurring over entire hydrocaulus, branches up to second order, in radiate planes; hydranth pedicels arising from main stem or branches of ¢rst order. Perisarc of main stem thick, ¢rm, single tubes 0.2^ 0.25 mm in diameter (Figures 2B & 3B), *0.02 mm, without annulations but with deformations. Branches with 1^2 rings at origin, 0.13^0.22 mm in diameter. Pedicels obscurely annulated at origin, with 3^8 rings, with thick perisarcal walls at origin, otherwise completely smooth throughout, 2.0^3.5 mm in length, *0.15 mm in diameter. Hydranths 0.6^0.7 mm in height, 0.5^0.6 mm in diameter (measured in the body region just below the tentacles), with a distinct groove at aboral region; hypostome large and £ared, 0.1^0.2 mm in height, 0.3^0.4 mm in diameter (Figures 1B, 2A & 3A); tentacles 27^30 in number, occurring in a whorl below hypostome. Nematocyst distribution on the tentacles giving them a spiny aspect (Figures 2G & 3E). General shape of hydranth vasiform, widening from half part of body to the whorl of origin of tentacles. Gonophores styloids, arising from body of a hydranth, immature placed in a circle around body of hydranth. Sex female, each with a simple spadix curving over a single egg. Tentacles of fertile hydranths may be reduced in number, to 18^22 (Figure 2D). Immature eggs almost circular, 6^7 per hydranth, *0.7 mm in diameter (Figures 1C, 2E & 3D). Perisarc covered pedicel of blastostyle conical, entirely corrugated, *0.9 mm in length (Figure 2D). Male gonophores not observed. Living and ¢xed hydranth body, hypostome, tentacles, coenosarc, and spadix deep blue due to abundant granules of pigment located in the gastrodermis (Figures 1 & 3). Nematocysts of one category, heterotrichous microbasic euryteles in two size-classes. Small microbasic euryteles (not seen discharged), 7.02.5 m, L/W¼2.8:1, oval, abundant; distributed over hydranth body, hypostome, coenosarc, regularly on tentacles (Figures 2G & 3E). Large microbasic euryteles (seen discharged), 19^23 12^15 mm, L/W¼1.53^1.58:1, bean-shaped, shaft crossing about 2/3 of the whole length of undischarged capsule, discharged capsule 219 mm, discharged shaft 13^15 mm in length, proportion S/C¼0.61^0.71; nematocysts abundant, distributed over hydranth body, hypostome, coenosarc, female spadix (Figure 2A & F). Etymology
This species is named in honour of Professor Klaus Ru«tzler (Smithsonian Institution, Washington, DC). Journal of the Marine Biological Association of the United Kingdom (2005)
DISCUSSION The worldwide distributed genus Eudendrium is one of the richest among anthoathecate hydroids in terms of number of species. The taxonomy of the group was based largely on gross morphology, which did not allow reliable identi¢cation of species. The use of the type, size and distribution of nematocysts as a taxonomic tool improved the situation considerably (Picard, 1951). Nowadays, the cnidome of about 50 species have been described and it is essential to use both morphology and nematocyst information for the taxonomy of the genus (Marques et al., 2000a). Eudendrium klausi sp. nov. is the only species of Eudendrium provided with pigment granules (Figures 1 & 3). These pigments, packed into endodermal cells, are abundantly distributed on the hydranth body, hypostome, tentacles, female spadix, and coenosarc. Other diagnostic characteristics of E. klausi sp. nov. are the gnarled stems, lacking the usual evenly parallel walls, the variable thickness of the stem perisarc (Figures 2B & 3B), and the conical pedicel of female gonophores (Figure 2D). Further di¡erences between species presenting a similar cnidome to that of E. klausi sp. nov. are summarized in Table 1. The cnidome of E. klausi sp. nov. is characterized by microbasic euryteles of two di¡erent dimensions. The small microbasic euryteles are common to all known cnidomes of Eudendriidae, therefore taxonomically not useful. On the other hand, the larger microbasic euryteles, distributed both on the hydranth body and/or hypostome, are shared by 15 nominal species, viz Eudendrium angustum Warren, 1908, Eudendrium arbuscula Wright, 1859, Eudendrium bathyalis Marques & Calder, 2000, Eudendrium caraiuru Marques & Oliveira, 2003, Eudendrium eximium Allman, 1877, Eudendrium fruticosum Allman, 1877, Eudendrium generale von Lendenfeld, 1885, Eudendrium kirkpatricki Watson, 1985, Eudendrium merulum Watson, 1985, Eudendrium minutum Watson, 1985, Eudendrium moulouyensis Marques et al., 2000b, Eudendrium novaezelandiae Marktanner-Turneretscher, 1890, Eudendrium pocaruquarum Marques, 1995, Eudendrium rameum (Pallas, 1766), Eudendrium ramosum (Linnaeus, 1758) (information on the cnidome of species from Millard, 1975; Watson, 1985; Marinopoulos, 1992; Marques, 1995, 2001; Schuchert, 1996; Marques & Calder, 2000; Marques et al., 2000a,b; Marques & Oliveira, 2003, and data from this study). Another species with similar cnidome is Eudendrium calceolatum Motz-Kossowska, 1905, but we concur with Marques et al. (2000b) that the species is of dubious validity. Although similar in cnidome, these 15 species of Eudendrium (including E. klausi sp. nov.) can be distinguished among each other on the basis of additional characters (see, for example, di¡erences between E. bathyalis, E. angustum, E. kirkpatricki, E. merulum, E. minutum, E. moulouyensis, E. novaezelandiae highlighted in Table 1). However, E. merulum is distinguished from other species on the basis of characteristics of the male gonophore, therefore not comparable to E. klausi sp. nov., here described from a female colony. Nonetheless, the species di¡er in the thicker hydrocauli and wider hypostome seen in E. klausi sp. nov., and the highly ornamented shaft present in the euryteles of E. merulum (cf. Watson, 1985; Bavestrello & Piraino, 1991; Marques, 2001).
Eudendrium klausi sp. nov. from Belize The distribution of the large microbasic euryteles is a distinguishing feature in some species of Eudendrium. For instance, E. arbuscula and E. caraiuru may be diagnosed by aggregations of nematocysts on the hydranth body, generally in pads (like E. caraiuru; cf. Marques & Oliveira, 2003) or, when extremely developed, forming a continuous ring (such as in E. arbuscula and exceptionally some hydranths of E. caraiuru as well; cf. Calder, 1972; Marques et al., 2000a; Marques & Oliveira, 2003). In the same way, abundant nematocysts on the hypostome are present in E. ramosum (Watson, 1985; Marques, 2001), di¡ering in this respect from E. klausi sp. nov. Some trophosome characters, although presumably variable (cf. Wedler, 1975), may also be useful to di¡erentiate Eudendrium species with a similar cnidome. However, trophosome characters are rarely unique for a given species. The lack of informative characters can lead to the so-called species complexes, like the ‘E. capillare’ species group (cf. Watson, 1985; Marques et al., 2000a) and the ‘E. glomeratum’ species group (cf. Marques & Oliveira, 2003). These complexes seem to be rather stable in their composition, therefore it appears justi¢ed to use characters of the trophosome in the taxonomy of the group. Nevertheless, E. klausi sp. nov. can be distinguished from other species (e.g. E. generale and E. rameum) on the basis of trophosome characters, such as its monosiphonic stem (cf. Millard, 1977; Watson, 1985; Marques et al., 2000a,b; Puce et al., 2002). Similarly, the entirely or extensively annulated hydrocauli of E. pocaruquarum (Marques, 1995) is also a trophosomal di¡erence to E. klausi sp. nov. Among the species of Eudendrium restricted to the Caribbean, three di¡er from E. klausi sp. nov. in their cnidome as they do not possess microbasic euryteles of two size-classes: E. capillare presents only small microbasic euryteles, E. bermudense has large macrobasic euryteles and, ¢nally, E. carneum has heterotrichous anisorhizas (besides its diagnostic female bi¢d spadix). The remaining seven Caribbean species of Eudendrium were only poorly described by Allman (1877) making their comparison with E. klausi sp. nov. di⁄cult or impossible. Allman (1877) did not describe any blue coloured species. Moreover, unlike E. klausi, Allman’s species were collected in deep waters. Anyhow, to complement our analysis, we re-studied these species, including a re-examination of available type-specimens. Of two out of these seven species, namely E. gracile and E. laxum, there exists no type material, and their descriptions are too vague for their establishment as valid species, therefore we regard them as species inquirenda. Five other species have type specimens deposited in the collections of the MCZ. The type material of E. attenuatum (MCZ 50237, colony without gonophores, USA, o¡ Florida, SSW Tortugas, BIBB Gulf Stream Exploration St. cast no. 3, 248290 4500 N 8381700000 W alcohol-formol preserved, 108 m, 18/01/1869, leg. L.F. de Pourtale's, det. G.J. Allman, holotype) consists of hydrocauli only, and apparently has only small microbasic euryteles. Indeed, Allman (1877:7) considered that in the absence, however, of hydranths and gonosome, the species is only provisionally referred to Eudendrium. The type material of E. tenellum (MCZ 50235, colony without gonophores, USA, o¡ Florida, o¡ Double-Head Journal of the Marine Biological Association of the United Kingdom (2005)
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Shot Key, Gulf Stream Expedition, 2385703000 N, 8082901500 W, alcohol preserved, 10/03/1869, 862 m, leg. L.F. de Pourtale's, det. G.J. Allman, holotype, without hydranths) also consists of hydrocauli without polyps, and apparently has only small microbasic euryteles. It is indeed remarkable that E. tenellum was quoted and redescribed so many times (e.g. Calder, 1972; Hirohito, 1988) based on the precarious description given by Allman (1877), which could correspond to any small Eudendrium colony. The material identi¢ed by Hirohito (1988) as E. tenellum Allman does evidently not belong to this species as the cnidome is distinctly di¡erent. Calder (1988), reviewing the status of several Allman’s nominal species, relegated E. tenellum to the status of a ‘nomen dubium’. Therefore, we also regard E. tenellum, in the sense of Allman (1877) as an indeterminate species. The holotype of E. exiguum (MCZ 35103, colony without gonophores, USA, Gulf Stream Exploration, o¡ Florida reef, alcohol preserved, 179 m, 06/05/1868, leg. F. de Pourtale's, det. G.J. Allman, holotype) has nematocysts other than microbasic euryteles, although they are di⁄cult to identify properly. Besides, the fragment of the type studied (2 mm wide and 2 cm long), is characterized by a heavy fasciculation, thus di¡erent from the general habit of E. klausi sp. nov. The two remaining species described by Allman (1877) are E. eximium (MCZ 35102, female colony, USA, Gulf Stream Expedition, Straits of Florida, west of Tortugas Island, cast no. 11, 24844015’’N 838460 W, Monroe Co., 16/ 01/1869, 79 m, col. L.F. de Pourtale's, det. G.J. Allman, alcohol, holotype) and E. fruticosum (MCZ 35105, male and female colonies, USA, Key West, FL, BIBB Florida Straits, o¡ Key West, Gulf Stream Expedition, 21/01/1869, 248180 0000 N, 8185001500 W, 247 m, col. L.F. Pourtale's, det. G.J. Allman, alcohol, holotype; see Figures 4A,B & 5A^ F), both comprise enough material, including fertile colonies. The data produced from the study of these species are presented in Table 1. Compared to E. klausi sp. nov., E. eximium di¡ers in the size of its nematocysts and E. fruticosum by the presence of a gelatinous pellicle enveloping the immature eggs. Besides, both E. eximium and E. fruticosum form fascicled colonies, and they do not have the unique characters found in E. klausi sp. nov., namely, the deep blue colour of hydranth, female spadix, and coenosarc. The colour pattern is an obvious taxonomic character in several invertebrate groups such as sponges, £atworms, molluscs, and arthropods. Hydroids are generally considered transparent, but several coloured structures such as red or black ocelli are often present in medusae. Generally, the colour of these structures is di⁄cult to observe in preserved materials because of the action of preserving liquids that makes the colour unsuitable for taxonomic purposes. Several species of hydroids are coloured through assimilation of pigments of their preys, such as in the red coloured Perarella schneideri which feed on the bryozoan Schizoporella longirostris (Bavestrello et al., 2000) or in the violet coloured Halocoryne epizoica which feed on the bryozoan Schizobrachiella sanguinea (Piraino et al., 1992). Many hydroid species, on the other hand, are coloured because of the presence of symbiotic microalgae (cf. Muscatine, 1974).
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Some species of Eudendrium are also coloured, especially when alive. For instance, E. moulouyensis is brown-green due to the presence of zooxanthellae (Marques et al., 2000b). Another species, E. generale, has Antarctic representatives described with an intense orange coloration, probably due to carotenoid pigments (Puce et al., 2002). A blue coloration, however, typical of the pelagic Porpita and Velella, is due to a carotenoprotein (Herring, 1971). Carotenoids are also responsible for the purple coloration of several hydrocoral species (Fox, 1972). This kind of pigmentation is not frequent among other hydroid species. The Mediterranean Synthecium evansi has blue colonies (G. Bavestrello, personal observation), while, in the case of Coryne eximia, the gonozooids only are coloured blue, making these polyps easily distinguishable from the gastrozooids found in the colony (Puce et al., 2003). Comparing the occurrence of blue coloration with established knowledge of hydrozoan systematics emphasizes the non-phylogenetic meaning of this characteristic. However, further investigation on the causes or nature of this unusual coloration may suggest some link between its expressions in di¡erent hydrozoan groups. We thank Alberto Lindner for suggestions and revision of the text. This work was ¢nancially supported by MIUR, Smithsonian Institution, FAPESP, and CNPq grants.
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Submitted 11 March 2004. Accepted 26 October 2004.
