Research Article |
Corresponding author: Tobias Lehmann ( lehmann@snsb.de ) Academic editor: Carsten Lüter
© 2014 Tobias Lehmann, Martin Heß, Roland R. Melzer.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lehmann T, Heß M, Melzer R (2014) Common littoral pycnogonids of the Mediterranean Sea. Zoosystematics and Evolution 90(2): 163-224. https://doi.org/10.3897/zse.90.7520
|
In the present study 21 littoral pycnogonid species from 5 families are analysed: Ammotheidae (9 species), Callipallenidae (5 species), Endeidae (2 species), Phoxichilidiidae (3 species), and Pycnogonidae (2 species). The material was mainly taken from Mediterranean pycnogonid collections housed in the Bavarian State Collection of Zoology. Additional material was collected during field trips. Altogether the material was obtained from six different locations: Banyuls-sur-Mer (France), Tavolara Island (Italy), Elba Island (Italy), Giglio Island (Italy), Sicily Island (Italy), and Istria Peninsula (Croatia). The animals were studied in detail with a scanning electron microscope (SEM). Additionally series of light microscopic pictures were made to generate extended depth of field pictures of whole animals. The observed features are compared with previous literature.
SEM, Pycnogonida , Sea spider, Achelia , Ammothella , Anoplodactylus , Callipallene , Endeis , Pycnogonum , Tanystylum , Trygaeus
The Pycnogonida or sea spiders are exclusively marine invertebrates, numbering more than 1330 species worldwide (
The phylogenetic position of the Pycnogonida has long been controversial and is still under debate. Today pycnogonids are placed either within the Chelicerata as sister taxon of the Euchelicerata or as sister taxon of all other Euarthropoda (
The Mediterranean Sea as we know it today developed about 5 million years ago. After the Messinian salinity crisis (5.6 to 5.3 Mya) the Mediterranean Sea was filled in a major flood approximately 5.3 million years ago, in which water poured in from the Atlantic Ocean and through the Strait of Gibraltar (
In the Mediterranean Sea 56 pycnogonid species are known until now. A current species list of the Mediterranean pycnogonid fauna is found in
Mediterranean pycnogonid literature is scattered (e.g.
There is a rapidly growing body of molecular studies especially on the Antarctic and Subantractic fauna (e.g.
In the present study representatives of some of the major genera of Mediterranean pycnogonids were sourced from the Bavarian State Collection of Zoology. Additional material was collected during field trips. A significant objective of this study is to remove any ambiguity in species identifications by providing a pictorial atlas principally based on high resolution Scanning Electron Microscope (SEM) images. Classification follows
Material: The majority of specimens were sourced from the Bavarian State Collection of Zoology. Additional material was collected during field trips to Banyuls-sur-Mer, France (June/July 2006) and Rovinj, Croatia (September 2006 and May 2007). Collecting sites are summarized in Figure
Light microscopy: Light microscopic pictures were taken using an Olympus SZX stereo microscope and a Jenoptic Prog-Res C12 digital camera (2580 × 1944 px; 96 dpi; colour depth 24 bit). Larger specimens (Endeis charybdaea, E. spinosa) were photographed using a Wild-Heerbrugg M5A stereo microscope and a Canon Digital IXUS 850 IS digital camera (3072 × 2304 px; 180 dpi; colour depth 24 bit). Up to 12 pictures with different focus steps along the z-axis were combined to a single respective image with a greater field of depth using the computer software Auto Montage (Syncroscopy) or Helicon Focus (HeliconSoft).
Scanning electron microscopy: For SEM preparation, specimens were dehydrated in a graded acetone series (70%, 80%, 90%, 10 min. each, plus 3 × 100%, 20 min. each) and critical-point-dried in a Baltec CPD 030. Dried specimens were mounted on SEM stubs with self-adhesive carbon stickers and coated with gold on a Polaron Sputter Coater. SEM pictures (2048 × 1536 px; 72 dpi; colour depth 8 bit) were made with a LEO 1430VP at 10–20 kV. Scales were inserted using the measurement utility of the SEM.
Nomenclature: The present study follows the nomenclature in
ab | abdomen | mf | movable finger |
ac | auxiliary claw | mo | mouth |
an | anus | ne | neck |
ce | cephalon | o | oviger-article |
cf | chelifore | oc | ocular tubercle |
cg | cement gland | ov | oviger |
ch | chela | p | palp-article |
cl | claw | pa | palp |
cs | compound spine | pp | propodus |
cu | cutting lamina | pr | proboscis |
cx | coxa | pt | protuberance |
ff | fixed finger | sb | segment border |
fm | femur | se | segment |
go | genital openings | so | slit organ |
ha | hair | sp | spine |
hs | hollow spine | tb | tibia |
le | leg | te | teeth |
lp | lateral process | tr | trunk |
ls | lateral sense organ | ts | tarsus |
This study is based on 21 Mediterranean species. In all cases, the major morphological characteristics correspond with published descriptions; exceptions see remarks section of each species.
Order Pantopoda Gerstäcker, 1863
Suborder Eupantopodida Fry, 1978
Superfamily Ascorhynchoidea Pocock, 1904
Family Ammotheidae Dohrn, 1881
Achelia echinata Hodge, 1864
Achelia langi (Dohrn, 1881)
Achelia vulgaris (Costa, 1861)
Ammothella appendiculata (Dohrn, 1881)
Ammothella biunguiculata (Dohrn, 1881)
Ammothella longipes (Hodge, 1864)
Ammothella uniunguiculata (Dohrn, 1881)
Tanystylum conirostre (Dohrn, 1881)
Trygaeus communis Dohrn, 1881
Superfamily Nymphonoidea Pocock, 1904
Family Callipallenidae Hilton, 1942
Callipallene emaciata (Dohrn, 1881)
Callipallene phantoma (Dohrn, 1881)
Callipallene producta (Sars, 1888)
Callipallene spectrum (Dohrn, 1881)
Callipallene tiberi (Dohrn, 1881)
Superfamily Phoxichilidoidea Sars, 1891
Family Endeidae Norman, 1908
Endeis charybdaea (Dohrn, 1881)
Endeis spinosa (Montagu, 1808)
Family Phoxichilidiidae Sars, 1891
Anoplodactylus angulatus (Dohrn, 1881)
Anoplodactylus petiolatus (Krøyer, 1844)
Anoplodactylus pygmaeus (Hodge, 1864)
Superfamily Pycnogonoidea Pocock, 1904
Family Pycnogonidae Wilson, 1878
Pycnogonum nodulosum Dohrn, 1881
Pycnogonum (Retroviger) pusillum Dohrn, 1881
Ammothea brevipes Hodge, 1864
Ammothea echinata Hodge, 1864
Ammothea fibulifera (Dohrn, 1881)
ZSMA20071461: male; Giglio Island, Italy; 08.04.2005; 4–8 m; brown algae. ZSMA20071462: female; Giglio Island, Italy; 11.04.2005; 20 m. ZSMA20071463: female; Rovinj, Croatia; 05.07.2005; 0–1.5 m; Stypocaulon scoparium (Linnaeus) Kützing, 1843.
Ammothea langi Dohrn, 1881
ZSMA20071464: male; Cape Savudrija, Croatia; 08.10.2004: 0–1 m; under stones. ZSMA20071465: male; Cape Savudrija, Croatia; 22.05.2005: 0–1 m. ZSMA20071466: female; Cape Savudrija, Croatia; 08.08.2005; 0–1 m; on stone.
According to
Achelia franciscana (Dohrn, 1881)
Alcinous vulgaris Costa, 1861
Ammothea fransciscana Dohrn, 1881
ZSMA20071467: male; Bayuls-sur-Mer, France; 07.2004. ZSMA20071468: female; Giglio Island, Italy; 08.04.2005; 8–13 m; brown algae. ZSMA20071469: female; Giglio Island, Italy; 08.04.2005; 8–13 m; brown algae.
Adult males are differentiated by the character described by
Here, only coxa 2 of the first leg has three protuberances with spine on the one and two on the other side; the coxa 2 of the other legs has two protuberances with spine on each side (Fig.
Ammothea appendiculata Dohrn, 1881
ZSMA20071470: female; Cape Savudrija, Croatia; 03.09.2004: 0–1 m. ZSMA20071471: female; Cape Savudrija, Croatia; 08.10.2004: 0–1 m; on stone. ZSMA20071472: female; Cape Savudrija, Croatia; 08.08.2005; 0–1 m; under stone.
Ammothea biunguiculata Dohrn, 1881
ZSMA20071473: male; Cape Savudrija, Croatia; 08.10.2004: 0–1 m; on stone.
Achelia longipes (Hodge, 1864)
Achelia magnirostris (Dohrn, 1881)
Ammothea longipes Hodge, 1864
Ammothella magnirostris (Dohrn, 1881)
ZSMA20071477: female; Bayuls-sur-Mer, France; 07.2004.
Ammothea uniunguiculata Dohrn, 1881
ZSMA20140003: male; Sicily, Italy; 1988
Clotenia conirostris Dohrn, 1881
ZSMA20071505: female; Bayuls-sur-Mer, France; 07.2004. ZSMA20071506: female; Bayuls-sur-Mer, France; 07.2004. ZSMA20071507: male; Bayuls-sur-Mer, France; 07.2004. ZSMA20071508: male; Bayuls-sur-Mer, France; 07.2004.
ZSMA20071509: female; Giglio Island, Italy; 12.04.2005. ZSMA20071510: male; Giglio Island, Italy; 12.04.2005. ZSMA20071511: male; Giglio Island, Italy; 12.04.2005. ZSMA20071512: male; Giglio Island, Italy; 12.04.2005. ZSMA20071513: male; Cape Kamenjak, Croatia; 03.05.2007; 0–5 m; Stypocaulon scoparium (Linnaeus) Kützing, 1843.
According to
Callipallene emaciata emaciata Stock, 1954
Callipallene emacinata (Dohrn, 1881)
Pallene emaciata Dohrn, 1881
ZSMA20071485: female; Giglio Island, Italy; 09.04.2005; 3–5 m; Halopteris sp.. ZSMA20071486: female; Giglio Island, Italy; 09.04.2005; 3–5 m; Halopteris sp.. ZSMA20071487: male; Giglio Island, Italy; 09.04.2005; 3–5 m; Halopteris sp..
Callipallene phantoma crinita Stock, 1952
Callipallene phantoma phantoma (Dohrn, 1881)
Pallene phantoma Dohrn, 1881
ZSMA20071489: female; Brucoli, Sicily, Italy; 14.09.1988. ZSMA20071490: female; Banyuls-sur-mer, France; 07.2004. ZSMA20071491: female; Giglio Island, Italy; 11.04.2005; 20 m; red algae. ZSMA20071492: male; Giglio Island, Italy; 11.04.2005; 20 m; red algae. ZSMA20071493: male; Giglio Island, Italy; 11.04.2005; 20 m; red algae. ZSMA20071494: male; Rovinj, Croatia; 05.07.2005.
Callipallene brevirostris producta (Sars, 1888)
Pallene producta Sars, 1888
ZSMA20071495: female; Giglio Island, Italy; 08.04.2005; 13–18 m; on brown algae. ZSMA20071496: female; Giglio Island, Italy; 08.04.2005; 13–18 m; on brown algae.
Callipallene emaciata spectrum (Dohrn, 1881)
Pallene spectrum Dohrn, 1881
ZSMA20140004: male; Banjole, Rovinj, Croatia; 07.1968; 5 m; on Cystoseira sp..
Callipallene emaciata tiberii (Dohrn, 1881)
Pallene tiberi Dohrn, 1881
ZSMA20071488: female; Giglio Island, Italy; 12.04.2005; 1 m; Halopteris sp..
Chilophoxus charybdaeus (Dohrn, 1881)
Phoxichilus charybdaeus Dohrn, 1881
ZSMA20071497: male; Rovinj, Croatia; 04.09.2006; 30 m.
Chilophoxus spinosus Montagu
Endeis gracilis Philippi, 1843
Endeis laevis (Grube, 1871)
Endeis vulgaris (Dohrn, 1881)
Phallangium spinosum Montagu, 1808
Phoxichilus vulgaris Dohrn, 1881
ZSMA20071498: female; Rovinj, Croatia; 04.2004. ZSMA20071499: male; Rovinj, Croatia; 05.05.2007; 0–2 m; Corallina elongata J.Ellis & Solander, 1786.
According to
Anoplodactylus angulirostris (Dohrn, 1881)
Phoxichilidium angulatum Dohrn, 1881
ZSMA20071478: female; Banyuls-sur-mer, France; 07.2004. ZSMA20071479: female; Giglio Island, Italy; 14.4.2005; 20 m; red algae.
Anoplodactylus guyanensis Child, 1977
Anoplodactylus hedgpethi Bacescu, 1959
Anoplodactylus longicollis (Dohrn, 1881)
Anoplodactylus pygmaeus (Hoek, 1881)
Pallene attenuata Hodge, 1864
Phoxichilidium attenuatus (Hodge, 1863)
Phoxichilidium longicolle Dohrn, 1881
Phoxichilidium petiolatum Krøyer, 1844
ZSMA20071480: male; Banyuls-sur-mer, France; 07.2004. ZSMA20042384: male; Bayuls-sur-Mer, France; 06.2003.
In literature there is diverse information about the articulation of the oviger, 7 articles in
Anoplodactylus derjugini (Losina-Losinsky, 1929)
Anoplodactylus exiguus (Dohrn, 1881)
Pallene pygmaea Hodge, 1864
Phoxichilidium exiguum Dohrn, 1881
Phoxichilidium pygmaeum Hoek, 1881
ZSMA20071481: female; Elba, Italy; 20.10.2004; 3–5 m. ZSMA20071482: female; Elba, Italy; 21.10.2004; 3–5 m. ZSMA20071483: male; Elba, Italy; 20.10.2004; 0.5–4 m. ZSMA20071484: male; Cape Kamenjak, Croatia; 03.05.2007; 0–5 m; Stypocaulon scoparium (Linnaeus) Kützing, 1843.
In the literature there is diverse information about the articulation of the oviger, 7 articles in
ZSMA20071503: female; Banyuls-sur-mer, France; 07.2004; 30 m; red algae. ZSMA20030112: female; Tavolara Island, Sardinia, Italy; 2002.
Pycnogonum pusillum Dohrn, 1881
ZSMA20071504: male; Brucoli, Sicily, Italy; 14.09.1988.
Achelia echinata, male; A: Dorsal view; scale 400 µm; B: Dorsal view of trunk, lateral processes touch each other; scale 200 µm; C: Frontal view of trunk, protuberances with spines on lateral process; scale 200 µm; D: Lateral view of proboscis, palp and chelifores with reduced chela; scale 100 µm; E: Mouth opening, dorsal is up; scale 20 µm; F: Mouth opening; scale 5 µm.
Achelia echinata, male; A: Chelifores with reduced chela; scale 40 µm; B: Distal articles of right 8-articled palp; scale 40 µm; C: Distal articles of 10-articled oviger; scale 40 µm; D, E: Compound spines on last oviger-article; scales 10 µm and 5 µm, respectively; F: Lateral process, coxa 1 and 2 of right 3rd leg, 2 protuberances with spine on each side of coxa 2; scale 100 µm; G: Coxa 2 with 2 protuberances with spine on each side (right 3rd leg); scale 40 µm.
Achelia echinata, male; A: Left 3rd leg; scale 200 µm; B: Lateral view of coxa 2 with genital protuberance (right 4th leg); scale 40 µm; C: Genital opening (right 3rd leg); scale 20 µm; D: Lateral view of femur with cement gland on distal part (left 3rd leg); scale 100 µm; E: Cement gland (right 4th leg); scale 40 µm; F: Tarsus, propodus, and claw, auxiliary claws about half as long as claw (left 3rd leg); scale 100 µm; G: Abdomen; scale 40 µm; H: Hair and slit organ on dorsal side of trunk; scale 5 µm.
Achelia echinata, female; A: Dorsal view; scale 400 µm; B: Ventral view of proboscis; scale 100 µm; C: Left 10-articled oviger; scale 100 µm; D: Lateral process, coxa 1 and 2 of right 3rd leg, lateral processes touch each other; scale 100 µm; E: Left 3rd leg; scale 200 µm; F: Ventral view of coxa 2 with genital opening, distal is right (left 3rd leg); scale 40 µm; G: Genital opening; scale 20 µm.
Achelia langi, male; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk; scale 400 µm; C: Ventral view of proboscis; scale 200 µm; D: Dorsal view of right lateral processes 1 and 2, 2 protuberances with spine on lateral process 1 and 1 on lateral process 2; scale 100 µm; E: Dorsal view of right lateral processes 3 and 4, 1 protuberance with spine on lateral process 3 and lateral process 4 without protuberance or spine; scale 100 µm; F: Left 3rd leg; scale 400 µm; insert: Genital protuberance; scale 100 µm.
Achelia langi, female; A: Dorsal view of trunk; scale 400 µm; B: Dorsal view of left lateral processes, 1 protuberance with spine on lateral process 1-3, lateral process 4 without protuberance or spine; scale 200 µm; C: Right 3rd leg; scale 400 µm; D: Ventral view of coxa 2 with genital opening, distal is down (right 3rd leg); scale 100 µm; E: Genital opening; scale 20 µm; F: Tarsus, propodus, and claw, auxiliary claws about 2/3 of length of claw; scale 100 µm; G: Abdomen with anus; scale 40 µm.
Achelia vulgaris, male; A: Dorsal view of trunk; scale 400 µm; B: Dorsal view of left lateral processes, lateral processes do not touch each other; scale 200 µm; C: 2 protuberances with spine on right side and 3 on left side of coxa 2, distal is down (right 1st leg); scale 100 µm; D: Lateral process, coxa 1 and 2, 2 protuberances with spine on each side of coxa 2 (left 3rd leg); scale 100 µm; E: Right 3rd leg; scale 400 µm; F: Tarsus, propodus, and claw, auxiliary claws about half as long as claw; scale 100 µm.
Achelia vulgaris, female; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk, lateral processes do not touch each other; scale 300 µm; C: Ventral view of proboscis; scale 100 µm; D: Right 10-articled oviger; scale 100 µm; E: Right 3rd leg; scale 400 µm; F: Dorsal view of coxa 2 (left 3rd leg); scale 40 µm; G: Ventral view of coxa 2 with genital opening (left 3rd leg); scale 40 µm.
Ammothella appendiculata, female; A: Dorsal view of trunk; scale 400 µm; B: Lateral view of trunk; scale 200 µm; C: Ventral view of trunk; scale 200 µm; D: Mouth opening, dorsal is up; scale 20 µm; E: Right chelifore with reduced chela and right 9-articled palp; scale 100 µm; F: Ocular tubercle with lateral sense organ; scale 40 µm.
Ammothella appendiculata, female; A: Lateral process of left 3rd leg; scale 40 µm; B: Coxa 1 of right 3rd leg with hollow spines; scale 40 µm; C: Hollow spine; scale 5 µm; D: Right 4th leg; scale 400 µm; E: Tarsus, propodus, and claws (right 4th leg); scale 100 µm; F: Hairs and slit organs of dorsal side of trunk; scale 10 µm.
Ammothella biunguiculata, male; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk; scale 400 µm; C: Lateral view of trunk, oviger dissected; scale 400 µm; D: Dorsal view of proboscis; scale 100 µm; E: Right chelifore with reduced chela; scale 40 µm; F: Right 9-articled palp; scale 100 µm.
Ammothella biunguiculata, male; A: Lateral view of ocular tubercle; scale 100 µm; B: Right 3rd leg; scale 400 µm; C: Ventral view of coxa 2 with genital opening, distal is left (right 3rd leg); scale 40 µm; D: Tarsus and propodus with reduced claw and dominating auxiliary claws; scale 100 µm; E: Reduced claw and dominating auxiliary claws; scale 20 µm; F: Dorsal view of trunk segment 4 and abdomen; scale 100 µm.
Ammothella longipes, female; A: Dorsal view of trunk; scale 200 µm; B: Lateral view of trunk, one protuberance without spine aside on lateral processes; scale 200 µm; C: Ocular tubercle; scale 100 µm; D: Right lateral process; scale 100 µm; E: Left 3rd leg; scale 400 µm; F: Tarsus, propodus, and claws (left 3rd leg); scale 100 µm.
Ammothella uniunguiculata, male; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk; scale 100 µm; C: Cephalon with proboscis, chelifore, and palps; scale 200 µm; D: Right 3rd leg; scale 200 µm; E: Tibia 2, tarsus, and propodus with claw (right 3rd leg); scale 100 µm; F: Propodus with claw; scale 20 µm.
Tanystylum conirostre, male; A: Distal articles of right oviger with spines on tip; scale 10 µm; B: Left 3rd leg; scale 100 µm; C: Lateral view of coxa 2 with genital opening on a protuberance, distal is right (left 4th leg); scale 40 µm; D: Genital opening; scale 10 µm; E: Cement gland on femur, distal is up (left 3rd leg); scale 20 µm; F: Tarsus, propodus, and claws (left 3rd leg); scale 100 µm; G: Hairs and slit organs on dorsal side of trunk; scale 10 µm.
Tanystylum conirostre, female; A: Dorsal view of trunk; scale 400 µm; B: Frontal view of trunk; scale 200 µm; C: Left 10-articled oviger; scale 40 µm; D: Distal articles of left oviger; scale 40 µm; E: Ventral view of coxa 2 with genital opening, distal is right (right 4th leg); scale 40 µm; F: Genital opening; scale 10 µm; G: Abdomen; scale 40 µm.
Trygaeus communis, male; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk; scale 400 µm; C: Lateral view of trunk with protuberances on midline; scale 400 µm; D: Mouth opening; scale 40 µm; E: Dorsal view of proboscis and chelifore without chela; scale 100 µm; F: Tip of chelifore without chela; scale 20 µm.
Trygaeus communis, male; A: Lateral view of proboscis and 7-articled right palp; scale 100 µm; B: Ocular tubercle with lateral sense organ; scale 40 µm; C: Right 9-articled oviger; scale 100 µm; D: Distal articles of right oviger; scale 40 µm; E: Protuberance on midline of segment 2; scale 40 µm; F: Right 3rd leg; scale 200 µm.
Trygaeus communis, male; A: Dorsal view of coxa 1 with protuberances on distal corners (right 3rd leg); scale 100 µm; B: Ventral view of coxa 2 with genital opening, distal is down (left 3rd leg); scale 40 µm; C: Genital opening; scale 10 µm; D: Cement gland on femur, distal is left (left 3rd leg); scale 40 µm; E: Tarsus, propodus, and claws (right 3rd leg); scale 100 µm; F: Abdomen with protuberance on tip; scale 100 µm; G: Hairs and slit organs on dorsal side of trunk; scale 10 µm.
Trygaeus communis, female; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk; scale 300 µm; C: Lateral view of trunk with protuberances on midline, frontal is right; scale 100 µm; insert: Protuberance on segment 2; scale 20 µm; D: Distal article of oviger; scale 40 µm; E: Dorsal view of coxa 1 without protuberances on distal corners (right 2nd leg); scale 40 µm; F: Ventral view of coxa 2 with genital opening, distal is down (right 3rd leg); scale 100 µm; G: Genital opening; scale 20 µm.
Callipallene emaciata, male; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk; scale 200 µm; C: Dorsal view of cephalon with rather slender neck; scale 100 µm; D: Right 3rd leg; scale 100 µm; E: Tarsus, strongly curved propodus, and claw, auxiliary claws about half as long as claw (right 3rd leg); scale 100 µm; F: Segment 3, 4 and abdomen; scale 40 µm.
Callipallene emaciata, female; A: Dorsal view; scale 1 mm; B: Frontal view; scale 1 mm; C: Dorsal view of trunk; scale 200 µm; D: Dorsal view of cephalon with rather slender neck; scale 200 µm; E: Left chela, dorsal is up; scale 40 µm; F: Tarsus, strongly curved propodus, and claw, auxiliary claws about half as long as claw (left 3rd leg); scale 100 µm.
Callipallene phantoma, male; A: Dorsal view of trunk; scale 200 µm; B: Ventral view of trunk; scale 200 µm; C: Dorsal view of cephalon with very long and slender neck; scale 100 µm; D: Ventral view of proboscis; scale 100 µm; E: Left chela, dorsal is down; scale 40 µm; F: Left 10-articled oviger; scale 100 µm.
Callipallene phantoma, male; A: Distal articles of left oviger with compound spines; scale 40 µm; B: Compound spines on last oviger-article; scale 5 µm; C: Left 3rd leg; scale 200 µm; D: Ventral view of coxa 2 with genital opening, distal is down (right 2nd leg); scale 20 µm; E: Genital opening; scale 5 µm; F: Tarsus, straight propodus, and claw with very short auxiliary claws (right 3rd leg); scale 40 µm; G: Hair and slit organ on dorsal side of trunk; scale 5 µm.
Callipallene phantoma, female; A: Dorsal view of trunk; scale 200 µm; B: Ventral view of trunk; scale 200 µm; C: Dorsal view of cephalon with very long and slender neck; scale 200 µm; D: Frontal view of cephalon; scale 100 µm; E: Mouth opening; scale 20 µm; F: Left chela, dorsal is up; scale 40 µm.
Callipallene phantoma, female; A: Right 10-articled oviger; scale 100 µm; B: Distalmost oviger-articles with compound spines; scale 20 µm; C: Right 3rd leg; scale 200 µm; D: Ventral view of coxa 2 with genital opening, distal is down (left 4th leg); scale 40 µm; E: Genital opening; scale 10 µm; F: Tarsus, straight propodus, and claw with very short auxiliary claws (left 3rd leg); scale 40 µm; G: Small auxiliary claws; scale 10 µm.
Callipallene producta, female; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk; scale 200 µm; C: Ventral view of trunk; scale 200 µm; D: Dorsal view of cephalon with very long and slender neck; scale 200 µm; E: Ventral view of proboscis; scale 100 µm; F: Ocular tubercle with lateral sense organ; scale 40 µm.
Callipallene producta, female; A: Distal article of right 10-articled oviger; scale 40 µm; B: Compound spines on oviger-article 7; scale 10 µm; C: Right 3rd leg; scale 400 µm; D: Ventral view of coxa 2 with genital opening, distal is down (left 3rd leg); scale 40 µm; E: Genital opening; scale 10 µm; F: Tarsus, slightly curved propodus, and claw with very long auxiliary claws (right 3rd leg); scale 100 µm; G: Claw and auxiliary claws; scale 40 µm.
Callipallene spectrum, male; A: Dorsal view of trunk; scale 100 µm; B: Lateral view of trunk; scale 100 µm; C: Dorsal view of cephalon; scale 100 µm; D: Cephalon with proboscis and chelifore; scale 100 µm; E: Left 3rd leg; scale 200 µm; F: Tibia2, tarsus, propodus, and claws (left 3rd leg); scale 100 µm.
Callipallene tiberi, female; A: Dorsal view of trunk; scale 200 µm; B: Dorsal view of cephalon with very short neck; scale 100 µm; C: Mouth opening; scale 20 µm; D: Left 3rd leg; scale 200 µm; E: Tarsus, strongly curved propodus, and claw, auxiliary claws about half as long as claw (left 3rd leg); scale 100 µm; F: Segment 3, 4 and abdomen; scale 40 µm.
Endeis charybdaea, male; A: Right lateral process 1 with 2 protuberances; scale 100 µm; B: Coxa 1 of right 1st leg with 2 protuberances; scale 100 µm; C: Right 3rd leg; scale 1 mm; D: Femur with 8 of overall 24 cement glands (left 3rd leg); scale 100 µm; E: Single cement gland; scale 5 µm; F: Tarsus, propodus, and claws (right 3rd leg); scale 200 µm; G: Abdomen; scale 200 µm.
Endeis spinosa, male; A: Dorsal view, abdomen lost due to preparation; scale 1 mm; B: Dorsal view of trunk; scale 400 µm; C: Dorsal view of proboscis; scale 200 µm; D: Tip of proboscis with few spines around mouth; scale 100 µm; E: Mouth opening; scale 20 µm; F: Cephalon with small protuberances with spine and without chelifores; scale 40 µm.
Endeis spinosa, male; A: Ocular tubercle; scale 100 µm; B: 7-articled oviger; scale 200 µm; C: Small spine on oviger-article 6; scale 10 µm; D: Right lateral process 1 with 2 protuberances; scale 100 µm; E: Right lateral process 3 with 1 protuberance; scale 100 µm; F: Coxa 1 of right 1st leg with 2 protuberances; scale 100 µm; G: Coxa 1 of right 3rd leg with 1 protuberance; scale 100 µm; H: Left 3rd leg; scale 400 µm.
Endeis spinosa, male; A: Lateral view of coxa 2 with genital opening on a protuberance (right 3rd leg); scale 100 µm; B: Genital opening; scale 20 µm; C: Femur with overall about 11 cement glands (left 3rd leg); scale 200 µm; D: Single cement gland; scale 10 µm; E: Tarsus, propodus, and claws (left 3rd leg); scale 200 µm; F: Spine, hair and slit organ on dorsal side of trunk; scale 10 µm.
Endeis spinosa, female; A: Dorsal view of trunk; scale 1 mm; B: Ocular tubercle; scale 100 µm; C: Right lateral process 3 without protuberance; scale 100 µm; D: Coxa 1 of right 3rd leg without protuberance; scale 100 µm; E: Right 3rd leg; scale 1 mm; F: Genital opening on ventral side of coxa 2; scale 40 µm; G: Abdomen; scale 200 µm.
Anoplodactylus angulatus, female; A: Ventral view of proboscis with conspicuous angles at corners (arrowheads); scale 40 µm; B: Ocular tubercle; scale 40 µm; C: Right lateral process 3 with spine; scale 40 µm; D: Right 3rd leg; scale 200 µm; E: Tarsus and propodus with hollow spine on propodus (right 3rd leg); scale 40 µm; F: Hollow spine on propodus with slit on tip (arrowhead); scale 5 µm.
Anoplodactylus petiolatus, male; A: Dorsal view of trunk; scale 200 µm; B: Lateral view of cephalon overreaching posterior part of proboscis, oviger dissected; scale 200 µm; C: Chelifores with developed chela; scale 40 µm; D: Left lateral process 3 with protuberance; scale 40 µm; E: Tarsus, propodus with cutting lamina, and claws (left 3rd leg); scale 40 µm; F: Cutting lamina on propodus (right 1st leg); scale 20 µm.
Anoplodactylus pygmaeus, male; A: Dorsal view; scale 400 µm; B: Lateral view of trunk; scale 200 µm; C: Dorsal view of trunk; scale 200 µm; D: Ventral view of trunk; scale 200 µm; E: Lateral view of cephalon overreaching posterior part of proboscis, proboscis with constriction encircling it subterminally (arrowheads); scale 100 µm; F: Ventral view of proboscis with constriction encircling it subterminally (arrowheads); scale 40 µm.
Anoplodactylus pygmaeus, male; A: Mouth opening; scale 20 µm; B: Left chelifore with developed chela; scale 20 µm; C: Ventral view of right palp, reduced to a small tubercle; scale 40 µm; D: Ocular tubercle with lateral sense organ; scale 40 µm; E: 6-articled oviger; scale 100 µm; F: Distal articles of left oviger; scale 20 µm.
Anoplodactylus pygmaeus, male; A: Right lateral process 3 with protuberance and spine; scale 20 µm; B: Left 3rd leg; scale 200 µm; C: Ventral view of coxa 2 with genital opening, distal is down (left 4th leg); scale 20 µm; D: Dorsal view of femur with cement gland, distal is down (right 4th leg); scale 20 µm; E: Tarsus, propodus with cutting lamina, and claws (left 3rd leg); scale 100 µm; F: Cutting lamina on propodus; scale 100 µm; insert: Detail of cutting lamina; scale 20 µm; G: Hairs and slit organs on dorsal side of trunk; scale 10 µm.
Anoplodactylus pygmaeus, female; A: Dorsal view of trunk; scale 200 µm; B: Ventral view of trunk, proboscis with a constriction encircling it subterminally (arrowheads); scale 200 µm; C: Right lateral process 3 with protuberance and spine; scale 40 µm; D: Ventral view of coxa 2 with genital opening (right 4th leg); scale 40 µm; E: Genital opening; scale 10 µm; F: Tarsus, propodus with cutting lamina, and claws (left 1st leg); scale 40 µm; G: Cutting lamina on propodus; scale 5 µm.
Pycnogonum nodulosum, female; A: Dorsal view; scale 1 mm; B: Dorsal view of trunk with protuberances on dorsal midline; scale 1 mm; C: Lateral view of trunk; scale 1 mm; D: Left 3rd leg with protuberances on femur; scale 400 µm; insert: Dorsal view of protuberances on femur; scale 200 µm; E: Tarsus, propodus, and claw without auxiliary claws (left 3rd leg); scale 100 µm; insert: Spines on propodus; scale 20 µm; F: Hair and slit organ on dorsal side of trunk; scale 5 µm.
Pycnogonum (Retroviger) pusillum, male; A: Dorsal view; scale 400 µm; B: Dorsal view of proboscis; scale 100 µm; C: Right 3rd leg; scale 100 µm; D: Tarsus, propodus, and claw without auxiliary claws (right 3rd leg); scale 40 µm; E: Abdomen; scale 100 µm; F: Protuberance, scattered over trunk and legs; scale 5 µm.
We thank Stefan Friedrich for expert technical assistance; Katharina Jörger and Roland Meyer for help with collecting the animals; and Stefan Füssl for help with preparing the figures in Adobe Photoshop.