Meliphagidae Vigors, 1825

Nguyen, Jacqueline M. T., Hand, Suzanne J. & Archer, Michael, 2016, The Late Cenozoic Passerine Avifauna from Rackham’s Roost Site, Riversleigh, Australia, Records of the Australian Museum 68 (5), pp. 201-230 : 204-209

Family Meliphagidae Vigors, 1825

Genus et species indet.

Figs 2–7

Material. AR21604, distal left humerus; QM F30825, distal right humerus; QM F30855, distal right ulna; AR17407, left carpometacarpus; QM F57899 (AR17401), right carpometacarpus; QM F22794, proximal left carpometacarpus; QM F30824, distal left tibiotarsus; QM F36374, proximal left tarsometatarsus; QM F57929 , distal left tarsometatarsus; QM F36648, distal right tarsometatarsus.

Measurements (mm). QM F22794: preserved length 9.2, proximal width 3.9, length of os metacarpale alulare 1.9. QM F30824: preserved length 6.6, distal width ca 2.3, depth of condylus lateralis ca 2.1, depth of condylus medialis ca 2.2. QM F30825: preserved length 8.9, distal width 4.4, depth of condylus dorsalis 2.2. QM F30855: preserved length 13.2, distal width (condylus dorsalis to tub. carpale) 3.2, depth of condylus dorsalis ca 2.8. QM F36374: preserved length 12.3, proximal width 2.7, proximal depth (lateral aspect)>2.0. QM F36648: preserved length 8.8, distal width 2.2, depth of tr II ca 1.0, depth of tr III 1.5, depth of tr IV 1.3. QM F57899 : preserved length 17.0, proximal width>3.9, length of os metacarpale alulare 2.0, distal width ca 3.5. QM F57929 : preserved length 11.0, distal width ca 2.0, depth of tr II ca 1.0, depth of tr III ca 1.5, depth of tr IV 1.2. AR17407: preserved length 10.6, proximal width>3.1, length of os metacarpale alulare>1.5, distal width>2.2. AR21604: preserved length 9.3, distal width>4.0, depth of condylus dorsalis 2.5.

Description and comparisons. Humerus. QM F30825 ( Fig. 2A,D ) is assigned to Meliphagidae because it shares the following suite of character states with this family. (1) The distal end is (1) greatly expanded dorsally and (2) ventrally from the shaft. (3) The sulcus humerotricipitalis is deep. (4) When viewed caudally, the distal profile of the humerus between the sulcus humerotricipitalis and sulcus scapulotricipitalis is very concave. (5) The proc. flexorius is truncate. (6) The proc. supracondylaris dorsalis is bifurcated and (7) projects far dorsally. (8) The fossa m. brachialis is moderately deep.

QM F30825 is slightly smaller than the humerus of the Bell Miner, Manorina melanophrys . It differs from other meliphagids examined in the following features. The distal end of QM F30825 is less expanded ventrally from the shaft than in Meliphaga , Anthochaera , Myzomela and Philemon , and less expanded dorsally than in Stomiopera and Melithreptus . The fossil further differs from Meliphaga , Stomiopera , Melithreptus and from Acanthorhynchus , Manorina and Nesoptilotis in having a deeper sulcus humerotricipitalis. The fossa m. brachialis is deeper in the fossil than in Philemon and Anthochaera . In QM F30825, the ventral apex of the proc. supracondylaris dorsalis is further set off from the distal end than in Manorina .

AR21604 ( Fig. 2C,F ) is a distal humerus with breakage to the dorsal portion. It is tentatively assigned to Meliphagidae because it shares features 2–5 and 8, as described above. AR21604 is similar in size to the humerus of the Tawnycrowned Honeyeater, Gliciphila melanops . It differs from Nesoptilotis , Melithreptus , Acanthorhynchus and Manorina in having a deeper sulcus humerotricipitalis. The fossa m. brachialis is deeper in AR21604 than in Stomiopera , Manorina , Philemon and Anthochaera . AR21604 differs from QM F 30825 in being smaller in size, and in having a more ventrally expanded distal end, a shallower sulcus humerotricipitalis and a slightly deeper fossa m. brachialis.

Within the Australian passerine avifauna, Petroicidae and Cracticidae are also characterised by a distal humerus that is considerably expanded dorsally and ventrally from the shaft. AR21604 can be excluded from Petroicidae because the distal end is deeper relative to its width, the sulcus humerotricipitalis is deep and the tub. supracondylare ventrale is more developed. This fossil can also be excluded from Cracticidae because it is considerably smaller in size, has a deep fossa m. brachialis and has a less developed ridge that ventrally bounds the sulcus scapulotricipitalis.

Ulna. QM F30855 ( Fig. 3A–C ) is tentatively assigned to Meliphagidae because it shares the following character states with this family. The tub. carpale is cranio-caudally compressed, short relative to the condylus ventralis, and perpendicular to the long axis of the shaft. The depressio radialis is deep. The condylus dorsalis protrudes well caudally from the shaft and its proximal extent is far greater than that of the condylus ventralis. The papillae remigales caudales are low. The sulcus intercondylaris is caudally deep and its distal profile is a moderately deep notch.

The fossil ulna is similar in size to the corresponding bone of Lewin’s Honeyeater, Meliphaga lewinii ( Fig. 3D–F ). Within Meliphagidae, QM F 30855 differs from the ulnae of Gliciphila , Manorina , Lichmera , Nesoptilotis , Conopophila and Philemon in that the condylus dorsalis protrudes farther caudally of the shaft. It differs from Philemon in having a squarer, rather than rounded, tub. carpale. The sulcus intercondylaris is shallower in the fossil than in Myzomela , but deeper than in Philemon . QM F30855 is distinguished from Anthochaera by its less developed papillae remigales caudales.

QM F30855 can be distinguished from similarly mediumto small-sized birds from other Australian passerine families by the following features. The fossil ulna differs from those of Dasyornithidae , Psophodidae , Pachycephalidae , Monarchidae , Artamidae , Locustellidae and Turdidae in that the sulcus intercondylaris is deep caudally and distally. The fossil differs from Rhipiduridae in having low papillae remigales caudales and from Orthonychidae in having a cranio-caudally shallow tub. carpale. It differs from Pomatostomidae and Sturnidae because the condylus dorsalis is greater in proximal extent relative to the condylus ventralis. QM F30855 further differs from Sturnidae in having a comparatively short tub. carpale, and from Pomatostomidae in having a deeper depressio radialis.

Carpometacarpus. QM F57899 ( Fig. 4A–B ) is a carpometacarpus with damage to the trochlea carpalis ventralis, caudal section and distal end. This fossil specimen is referred to Meliphagidae because it exhibits the following combination of features. (1) The carpometacarpus is straight and elongate. (2) The distal edge of the facies articularis alularis is located proximally of the large fovea lig. ventalis. (3) The fovea carpalis caudalis is deep and its distal margin is situated distally of the proc. cranialis. (4) The proc. dentiformis is low and (5) arises at about the proximo-distal midpoint of the os metacarpale majus. (6) The spatium intermetacarpale is long and narrow. (7) In dorsal aspect, a small portion of the spatium intermetacarpale is visible proximally of the proc. intermetacarpalis. (8) The distal end of the os metacarpale majus is broad and its cranial extent is greater than that of the proc. dentiformis. (9) The distal end of the sulcus tendinosus broadens into a large fossa on the distal end of the os metacarpale majus. (10) There is a distinct ventral fossa on the distal end of the os metacarpale minus. QM F57899 is slightly smaller in size than the carpometacarpus of the Noisy Miner, Manorina melanocephala ( Fig. 4C–D ). It differs from the extant honeyeaters studied here by the following features. QM F57899 differs from all meliphagids studied except for Xanthotis , Meliphaga and Anthochaera in having a deeper fossa at the distal end of the sulcus tendinosus. It differs from the carpometacarpi of Stomiopera , Nesoptilotis , Philemon , Anthochaera and Lichmera in having a larger, more defined proc. dentiformis. It differs from Melithreptus , Gliciphila , Ptilotula , Myzomela , Acanthorhynchus and Manorina in possessing a deeper ventral fossa on the distal end of the os metacarpale minus. The fossil differs from Stomiopera , Anthochaera , Ptilotula and Meliphaga in having a larger, more convex proc. cranialis. It differs from Acanthorhynchus , Meliphaga , Myzomela and Lichmera in having a deep notch in the distal margin of the os metacarpale minus. It differs from Conopophila , Xanthotis , Melithreptus and Anthochaera in having a deeper fovea lig. ventralis. The fossil is similar in proportions and overall morphology to species of Manorina studied, but its fragmentary nature precludes confident identification to genus level.

QM F22794 ( Fig. 4E–F ) and AR17407 ( Fig. 4G–H ) are tentatively referred to Meliphagidae because they share the following features with this family. QM F22794 shares characters 1–3, 5 and 7 described above, whereas AR17407 exhibits characters 1, 3, 5, 7 and 10. QM F22794 is similar in size to the corresponding element of Meliphaga lewinii . It differs from Ptilotula , Conopophila , Xanthotis and Meliphaga in having a deeper fovea lig. ventralis. The fossil differs from Nesoptilotis , Gliciphila , Stomiopera and Ptilotula in having a larger and more convex proc. cranialis. AR17407 corresponds in size to the carpometacarpus of Gliciphila melanops ( Fig. 4I–J ). It differs from Stomiopera , Anthochaera and Xanthotis in having a shallower fovea carpalis caudalis. The fossil differs from Xanthotis , Conopophila and Myzomela in possessing a smaller proc. cranialis. It further differs from Xanthotis and Conopophila in having a deeper fovea lig. ventralis. The fossil is broken proximally of the apex of the proc. dentiformis but preserves the base of the process, where the os metacarpale majus becomes wider. This base suggests that the proc. dentiformis was more developed in AR17407 than in Stomiopera , Anthochaera , Philemon and Nesoptilotis , where it is very low. AR17407 differs from QM F22794 and QM F 57899 in its considerably smaller size, and in having a shallower fovea carpalis caudalis and a smaller proc. cranialis. QM F22794 is overall similar in size and morphology to QM F57899 ,

although the former only preserves the proximal end. The fragmentary preservation of these fossils precludes further determination of their relationships.

Apart from meliphagids, a distally situated fovea carpalis caudalis relative to the proc. cranialis is also present in campephagids, pachycephalids, corvids and monarchids. However, the tentatively assigned fossils can be excluded from these birds (except Colluricincla ) because they possess a proc. dentiformis. QM F22794 and AR17407 differ from species of Colluricincla examined in having a shorter fovea lig. ventralis and in lacking a groove cranially of the proc. pisiformis.

Tibiotarsus. QM F30824 ( Fig. 5A ) is provisionally assigned to Meliphagidae because it exhibits the following suite of character states. The pons supratendineus is very long relative to its width. The tuberositas retinaculi extensori lateralis is low, elongate and situated on the proximal edge of the pons supratendineus. The bony ridges for attachment of the retinaculum m. fibularis are low and long. The medial bony ridge is approximately level with the tuberositas retinaculi extensori lateralis.

QM F30824 is similar in size to the corresponding bone of the Yellow-tinted Honeyeater Ptilotula flavescens ( Fig. 5B ). It differs from the extant meliphagids examined in this study in the following features. The fossil tibiotarsus differs from the corresponding bone of Acanthorhynchus , Stomiopera , Anthochaera and Lichmera in having a relatively shorter pons supratendineus. It differs from Ptilotula and Melithreptus in that the distance between the tuberositas retinaculi extensori medialis and its lateral counterpart is smaller. The tuberositas retinaculi extensori lateralis is situated further proximally relative to the pons supratendineus in the fossil than in Myzomela and Conopophila .

The tibiotarsal morphologies of meliphagids and acanthizids are very similar but can be differentiated by the following combination of features. In species of meliphagids examined, the bony ridges for the retinaculum m. fibularis are less pronounced and relatively longer than in acanthizids. The sulcus m. fibularis is shallower and the incisura intercondylaris is slightly wider. The tuberositas retinaculi extensori lateralis is less prominent and is situated further proximally on the pons supratendineus than in acanthizids studied.A long pons supratendineus is a characteristic feature of meliphagids, acanthizids and pachycephalids, but QM F30824 can be distinguished from the latter by its relatively longer pons and more cranially located epicondylus medialis. Also, the tuberositas retinaculi extensori medialis is situated further proximally of the bony ridges for the retinaculum m. fibularis in the fossil than in pachycephalids examined.

Tarsometatarsus. QM F36374, QM F36648 and QM F57929 ( Figs 6–7 ) are referred to Meliphagidae because they possess the following combination of characters, including those identified by Boles (2005). The cotyla medialis is greater in proximal extent than the cotyla lateralis. In proximal view, the planto-medial corner of the proximal end protrudes further medially than the rim of the cotyla. The foramina vascularia proximalia are located distally of the arcus extensorius. The tuberositas m. tibialis cranialis is low and situated well distally of the arcus extensorius. This tuberosity is located medially of the medio-lateral midpoint of the shaft. When viewed medially, the impressio lig. collateralis medialis is situated distally of the level of the arcus extensorius. The medial depth of the shaft is moderately deep at about level with the tuberositas m. tibialis cranialis. The lateral edge of the sulcus extensorius is low and the dorsal shaft surface is subsequently near perpendicular to the medial and lateral surfaces. The distal end of the tarsometatarsus is narrow, dorso-plantarly compressed and bent plantarly from the shaft. The fossa metatarsi I is large and deep. The medial shaft edge expands medially at about level with the edge of the fossa metatarsi I to form a flange that appears very shallow in medial aspect. The trochlea metatarsi II is short, latero-medially compressed and has a pointed distal profile. It is rotated planto-medially from the long axis of the shaft, such that the medial facies and fovea lig. collateralis medialis of the trochlea metatarsi II are visible in dorsal aspect. The incisura intertrochlearis medialis is narrower than the incisura intertrochlearis lateralis. The medial rim of the trochlea metatarsi III is greater in dorsal and distal extent than the lateral rim. The lateral edge of the trochlea metatarsi IV is located medially of that of the shaft.

QM F36374 ( Fig. 6A ) is similar in size to the proximal tarsometatarsus of the White-eared Honeyeater, Nesoptilotis leucotis . This fossil is distinguished from the extant meliphagids studied here by the following character states. It differs from Melithreptus , Acanthorhynchus and Anthochaera in that the proximal end is wider relative to the shaft, and further differs from Anthochaera in having a shallower sulcus flexorius. The fossil differs from Melithreptus , Ptilotula , Lichmera , Gliciphila , Anthochaera in having a more elevated impressio lig. collateralis medialis. It also differs from Gliciphila and Anthochaera in having a shallower medial depth of the shaft.

QM F36648 ( Fig. 7B,E,H ) corresponds in size to the Bell Miner, Manorina melanophrys , whereas QM F57929 ( Fig. 7A,D,G ) is similar in size to N. leucotis ( Fig. 7C,F,I ). The fossils differ from the extant meliphagid species studied here in the following features. They differ from Myzomela , Philemon and Meliphaga in having a poorly developed medial flange at about level with the fossa metatarsi I. The fossils differ from Myzomela , Gliciphila , Conopophila and Anthochaera in lacking a second medial extension of the shaft at about level with the distal edge of the fossa metatarsi I. QM F36648 and QM F57929 differ from Nesoptilotis and Philemon in that the foramen vasculare distale is situated relatively further proximally from the incisura intertrochlearis lateralis. They differ from Lichmera , Acanthorhynchus and Manorina in that the lateral edge of the trochlea metatarsi IV is situated further medially of that of the shaft.

QM F57929 differs from all extant meliphagids examined except for Gliciphila and Conopophila in that the trochlea metatarsi II is not as medially inflected. It differs from Xanthotis , Nesoptilotis , Philemon , Melithreptus , Acanthorhynchus , Anthochaera and Manorina in that the medial rim of the trochlea metatarsi III does not extend as far distally relative to the lateral rim. QM F57929 differs from QM F 36648 in that the distal end is narrower relative to the shaft width, the foramen vasculare distale is situated relatively further proximally, and the medial flange at about level with the fossa metatarsi I is less developed. These differences suggest that QM F36648 and QM F57929 represent two different taxa.

Remarks. Meliphagids (honeyeaters) are a very large and diverse radiation of passerines endemic to the southwestern Pacific region, with about 178 species in 51 genera (Dickinson & Christidis, 2014). These birds are prominent elements of the Australian avifauna and act as important pollinators for many flowering plants ( Longmore, 1991). As well as nectar, honeyeaters feed on fruit, lerps and psyllid larvae ( Longmore, 1991; Schodde & Mason, 1999). Honeyeaters range in size from small to medium-large and utilise habitats ranging from rainforest to semi-arid woodland to subalpine shrubland ( Higgins et al., 2001).

Boles (2005) reported meliphagid tarsometatarsi from Riversleigh Faunal Zone C (middle Miocene) assemblages and from Rackham’s Roost Site. QM F36374 is similar in size to the fossils that preserve the proximal tarsometatarsus in Boles (2005), but it is not clear that it represents the same taxon. QM F36648 likely represents a different taxon from the fossil meliphagids in Boles (2005) because it is proportionately broader, larger in size and has a more prominent medial flange at about level with the fossa metatarsi I. QM F57929 may also represent a different taxon from the previously reported Riversleigh meliphagids because it is larger in size but slightly narrower, and has a more developed medial flange and a less medially inflected trochlea metatarsi II.

Fossils of honeyeaters have been recovered from several Quaternary sites in South Australia, Victoria and Western Australia ( Hope et al., 1977; Baird, 1991a). These fossils include a skull of Manorina melanocephala from the late Pleistocene Green Waterhole Cave in South Australia ( Baird, 1985), and material referred to cf. Gavicalis (Lichenostomus) virescens from Pleistocene cave deposits in the Nullarbor Plains region (Baird, 1990, 1991a). Holocene remains assigned to cf. Anthochaera carunculata and cf. A. chrysoptera from Amphitheatre Cave in Victoria were also reported ( Baird, 1992). Today, honeyeaters are abundant in the Riversleigh region, with 19 species having been recorded.

Figures

Figure 2. Humeri of(A,C,D,F) fossil Meliphagidae gen.et sp. indet., compared with the humerus of (B,E) Conopophila albogularis AM O.70096. (A,D) QM F30825, distal right humerus. (C,F) AR21604, distal left humerus. (A–C) caudal and (D–F) cranial aspects. Some characteristic features of Meliphagidae shown: 1, sulcus humerotricipitalis deep; 2, distal profile between sulci humerotricipitalis et scapulotricipitalis highly concave; 3, distal end greatly expanded dorsally and 4, ventrally;5, proc.supracondylaris dorsalis bifurcated and projects well dorsally;6, fossa m. brachialis moderately deep. Scale bar = 2 mm.

Figure 3. Distal ulnae of (A–C) fossil meliphagid gen. et sp. indet. QM F30855 and (D–F) Meliphaga lewinii AM O.60079 (mirrored). (A,D) cranial, (B,E) caudal and (C,F) dorsal aspects. Some characteristic features of Meliphagidae shown: 1, depressio radialis deep; 2, tub. carpale short and perpendicular to the shaft long axis; 3, condylus dorsalis far greater in proximal extent than condylusventralis;4, sulcus intercondylaris deep both caudally and distally; 5, papillae remigales caudales low; 6, condylus dorsalis protrudes well caudally from the shaft. Scale bar = 2 mm.

Figure 4. Carpometacarpi of fossil Meliphagidae gen. et sp. indet., compared with those of extant meliphagid species in (A,C,E,G,I) ventral and(B,D,F,H,J) dorsal aspects.(A–B) QM F57899, right carpometacarpus.(C–D) Manorina melanocephala AM O.59876. (E–F) QM F22794, proximal left carpometacarpus. (G–H) AR17407, left carpometacarpus. (I–J) Gliciphila melanops AM O.65515. Some characteristic features of Meliphagidae shown: 1, distal edge of facies articularis alularis situated proximally of the level of the fovea lig. ventralis; 2, fovea lig. ventralis large; 3, distal end of os metacarpale majus broad and its cranial extent is greater than that of proc. dentiformis; 4, fossa on ventral surface of distal end of os metacarpale minus; 5, fovea carpalis caudalis deep, its distal margin located distally of the level of proc. cranialis; 6, broad fossa at distal end of sulcus tendinosus. Scale bar = 2 mm.

Figure 5. Distal left tibiotarsi of (A) fossil meliphagid gen. et sp. indet. QM F30824 and (B) Ptilotula flavescens AM O.66333 in cranial aspect.Some characteristic features of Meliphagidae shown: 1, pons supratendineus very long; 2, tuberositas retinaculi extensori lateralis low and situated on proximal edge of pons supratendineus; 3, bony ridges for attachment of retinaculum m.fibularis low.Scale bar = 2 mm.

Figure 6. Proximal left tarsometatarsi of(A) fossil meliphagid gen. et sp. indet.QM F36374 and (B) Stomiopera unicolorAM O.70570 in dorsal aspect. Some distinguishing features for Meliphagidae shown: 1, cotyla medialis greater in proximal extent than cotyla lateralis; 2, impressio lig. collateralis medialis situated distally of arcus extensorius; 3, tuberositas m. tibialis cranialis located well distally of arcus extensorius and medially of the shaft midpoint. Scale bar = 2 mm.

Figure 7. Distal tarsometatarsi of(A–B,D–E,G–H) fossil Meliphagidae gen.etsp.indet., comparedwith that of (C,F,I)Nesoptilotis leucotis AM O.59863.(A,D,G)QM F57929,distal lefttarsometatarsus.(B,E,H) QM F36648, distal right tarsometatarsus. (A–C) dorsal and (D–F) plantar aspects, and (G–I) lateral aspect of distal end. Some characteristic features of Meliphagidae shown: 1, tr II medio-laterally compressed, pointed distally and rotated planto-medially from shaft long axis; 2, lateral edge of tr IV situated medially of that of shaft; 3, medial rim of tr III greater in distal extentthan lateral rim; 4, incisuraintertrochlearis medialis narrow; 5, medial flange at about level with fossa metatarsi I; 6, distal end dorso-plantarly compressed and distinctly bent plantarly. Scale bar = 2 mm.