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Fossil Legume Woods of the Prioria-Clade

Fossil Legume Woods of the Prioria-Clade

Review of Palaeobotany and Palynology 246 (2017) 44–61

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Review of Palaeobotany and Palynology

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Fossil legume of the -clade (subfamily ) from the lower Miocene (early to mid-Burdigalian) part of the Cucaracha Formation of Panama (Central America) and their systematic and palaeoecological implications

Oris Rodríguez-Reyes a,b,d,⁎, Peter Gasson c,HowardJ.Falcon-Langd, Margaret E. Collinson d a Smithsonian Tropical Research Institute, Box 0843-03092, Balboa, Ancon, Panama b Departamento de Botánica, Universidad de Panamá, Panama c Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, UK d Department of Earth Sciences, Royal Holloway, University of London, Egham, Surrey TW20 0EX, UK article info abstract

Article history: Three fossil specimens are described from the Miocene (early to mid-Burdigalian) part of the Cucaracha Received 26 August 2016 Formation of Panama, Central America. The calcareously-permineralised fossils, which contain Teredolites bor- Received in revised form 9 June 2017 ings, occur in erosive-based pebbly conglomerate lenses, interpreted as tidally-influenced fluvial channel de- Accepted 15 June 2017 posits. Detailed investigation of fossil wood anatomy reveals features characteristic of the Prioria-clade, a Available online 19 June 2017 supergenus of the legume subfamily, Detarioideae. Based on quantitative comparison with extant material in the micromorphology slide collection at the Royal Botanic Gardens, Kew, the fossil material is referred to two Keywords: new , Prioria hodgesii sp. nov. and Prioria canalensis sp. nov. Facies data imply that these new taxa may Panama isthmus have occupied a similar ecological niche to the extant Prioria , a saline-tolerant tropical that Inside Wood forms wetland gallery along tidal estuaries in Panama today. Findings contribute to the understanding Legume of the palaeoecology of this early-diverging subfamily within the basal Leguminosae. They, also, further extend Prioria knowledge of the coastal forests along the leading edge of North America during its Miocene convergence with South America. © 2016 Elsevier B.V. All rights reserved.

1. Introduction Detarioideae (84 genera) and a more derived and diverse clade com- prises Duparquetioideae (1 genus), Dialioideae (17 genera), Leguminosae (colloquially referred to as legumes; also validly (148 genera) and Papilionoideae (503 genera). This known as ) is the third most speciose family of flowering interpretation differs from the traditional view of a basal paraphyletic after the orchids (Orchidaceae) and daisies (Asteraceae/ caesalpinioid-grade from which arise the derived ‘Mimosoideae’ and Compositae) with ~765 genera and ~19,328 species (Sprent, 2001; ‘Papilionoideae’ clades (Lewis et al., 2005). Lewis et al., 2005; LPWG, 2017). Molecular evidence points to an In this paper, we describe new fossil woods from lower Miocene early Paleogene (~60 Ma) (Sprent, 2007) origin, with the oldest strata of Panama, Central America that may be confidently referred to well-accepted fossils of Eocene age (~52 Ma; Herendeen et al., the Leguminosae. While some legume genera show very distinctive 1992); the family now constitutes a major ecological component of wood anatomy (e.g., Robinia and Dalbergia in the Papilionoideae; the Earth's vegetation, including many taxa of economic importance Wheeler and Baas, 1992), others show a very generalised structure (LPWG, 2017). with many intergrading features (e.g., Lonchocarpus and , Based on comprehensive analyses of plastid matK gene sequences, a also in the Papilionoideae; Baretta-Kuipers, 1981; Gasson et al., new phylogeny has revealed six subfamilies in two clades (LPWG, 2004). As a result, the attribution of isolated fossil wood types to 2017): a basal clade comprises (12 genera) and the Leguminosae family can be very challenging indeed (Wheeler and Manchester, 2002). Nonetheless, the following constellation of features seen in our material is generally characteristic of legumes: ⁎ Corresponding author at: Smithsonian Tropical Research Institute, Box 0843-03092, – Balboa, Ancon, Panama. simple perforation plates, alternate intervessel pitting, vessel ray E-mail address: [email protected] (O. Rodríguez-Reyes). pits similar to intervessel pits in size and shape, fibres with simple

http://dx.doi.org/10.1016/j.revpalbo.2017.06.005 0034-6667/© 2016 Elsevier B.V. All rights reserved. O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 45 pits, vestured pits present (with some exceptions) and paratracheal understanding of the of the subfamily, and findings also axial parenchyma with mostly 2–4 cells per strand (Baretta-Kuipers, add to the wealth of new systematic and biogeographical knowledge 1981; Herendeen, 2000; Gasson et al., 2003). about the Miocene forests of Central America, prior to the formation The fossils reported here are of especial interest, however, and uplift of the Isthmus of Panama. because not only do they bear close comparison with legumes in general, but also they may be confidently referred to the early-di- verging Detarioideae clade, which is sister to almost all other 2. Geological context Leguminosae (LPWG, 2017) and mostly comprises large pan-tropical , distributed through , Asia and South America The fossil wood specimens, reported here, were collected from (Herendeen et al., 2003). This group underwent initial diversifica- Hodges Hill on the Gaillard Cut of the Panama Canal (Fig. 1), c. 25 km tion in Eocene times (De Franceschi and De Ploëg, 2003), with a sec- west of Panama City (Latitude 09°02′51.75″N; Longitude 79°39′14.02″ ondary diversification event commencing in early Miocene times W). Fossil woods are abundant at this important fossil locality, and com- (De la Estrella et al., 2017). The earliest fossil records of the prise calcareously-permineralised trunks, 0.31–3.0 m long and typically Detarioideae are probably -like pollen grains from the Eocene 0.25–0.80 m in diameter, commonly with Teredolites borings. The al- of North America (Muller, 1981, 1984); however, the oldest known lochthonous assemblage is preserved within a pebbly sandstone bed, fossils in Central America occur near the Oligocene–Miocene bound- interpreted as the deposits of a tidally-influenced fluvial channel ary (Hueber and Langenheim, 1986; Poinar and Poinar, 1999; Poinar (Rodríguez-Reyes et al., 2014). The bed containing the fossil wood as- and Chambers, 2015). The Detarioideae subfamily is particularly im- semblage is positioned ~20 m above the base of the Cucaracha Forma- portant because it is the subject of intense anatomical and molecular tion, that has been dated as early Miocene (early to mid-Burdigalian) investigation, for the information it might provide about the evolu- age (MacFadden et al., 2014). A welded silicic tuff (known as the tion and systematics of the Leguminosae family (Redden et al., “Cucaracha Tuff”), positioned stratigraphically ~50 m above the fossil 2010; LPWG, 2017). Our new fossils therefore contribute to the wood layer, has a U–Pb age of 18.81 ± 0.30 Ma and Ar/Ar age of 18.96

Fig. 1. Location details for the fossil site. 1., Panama in Central America. 2., The Panama Canal Zone showing the location of the Hodges Hill locality. Taken from Rodríguez-Reyes et al. (2014). 46 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61

±0.90Ma(MacFadden et al., 2014; Farris et al., 2017), providing a min- Archaeology, Smithsonian Tropical Research Institute, Panama. imum age for the fossil assemblage reported here (Fig. 2). Petrographic thin sections were prepared for each specimen in Trans- verse Section (TS), Radial Longitudinal Section (RLS) and Tangential 3. Materials and methods Longitudinal Section (TLS) in the Department of Earth Sciences, Royal Holloway University of London. Oriented fossil slices were mounted μ This paper builds on earlier descriptions of fossil wood from the on glass slides using EpoFix , ground to a thickness of ~30 m, fi Cucaracha Formation of Panama, including representatives of and cover slips were af xed with Canada balsam. Material was observed Malvaceae (Rodríguez-Reyes et al., 2014) and Malpighiales and imaged using an Olympus binocular BH-5 microscope with a Nikon (Rodríguez-Reyes et al., 2017) among a rich and diverse wood flora digital camera system and software. Modern woods in the Jodrell Labo- that also includes families such as Sapotaceae, Arecaceae, ?Meliaceae, ratory of the Royal Botanic Gardens, Kew, U.K. (RBGK) were also studied and other different types assigned to Malvaceae and Leguminosae for comparative purposes, and imaged using a Leica DM LB microscope (Rodríguez-Reyes, 2014). Anatomical preservation of fossil wood is gen- with Zeiss Axiocam HRc camera attachment and Zeiss Axiovision erally good, allowing for detailed description and comparison with mi- software. croscope slides of extant woods in reference collections (Fig. 2). 3.2. IAWA feature description 3.1. Accession data, specimen preparation, and imaging Specimens were described using the International Association of The three new fossil wood specimens, reported here, collected in Wood Anatomists (IAWA) List of Features for Hardwood Identification 2011 and 2013, are accessioned as STRI 14165, STRI 36273, and STRI (IAWA Committee, 1989). In this standard descriptive system, anatom- 36400 in the collections of the Center for Tropical Paleoecology and ical characters (termed features) are given a numerical code. Most

Fig. 2. Stratigraphic log of the Miocene Formation in the Panama Canal Basin, showing the horizon containing the permineralised woods, studied herein, within the Cucaracha Formation (after Montes et al., 2012). Taken from Rodríguez-Reyes et al. (2014). O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 47 features are qualitative (requiring determination of presence or ab- we broadly followed the strategies recommended by Wheeler (2011) sence) but others are quantitative (requiring measurement of a popula- and Falcon-Lang et al. (2012) and generally avoided putting too much tion). To obtain quantitative data for vessel and ray density, and the emphasis on variable quantitative features. Guided by the short-lists degree of vessel grouping, measurements were made in 10 different of taxa obtained from IWD searches, we compared the fossil woods fields of 1 mm2 area. For other quantitative features (e.g., mean vessel with the micromorphology slide collection of extant woods in the diameter, intervessel pit diameter, vessel–ray pit diameter, vessel ele- Jodrell Laboratory of the Royal Botanic Gardens, Kew, UK (RBGUK). ment length, ray height), a minimum of 25 measurements was obtain- We also examined key publications about the wood anatomy of partic- ed, but where preservation allowed, 50 measurements were obtained. ular groups. As part of our fossil descriptions, anatomical features are listed as a string of numerical IAWA codes. In this list, features may be qualified 4. Fossil wood description by one of two signs as follows: “?” indicates that there is uncertainty as to whether the feature is present and “v” indicates that the feature Order: Bromhead, 1838 is variable in occurrence. In addition to the normal IAWA descriptive Family: LEGUMINOSAE Jussieu, 1789 system, we also quantified the normal axial canal diameter because Subfamily: Detarioideae Burmeister, 1837 this was evidently a significant feature in distinguishing different ana- Supergenus: Prioria Greisbach 1860 (Breteler, 1999) tomical types. Discussion: The constellation of anatomical features seen in the three fossil wood specimens indicates an affinity with the 3.3. Inside Wood Database and comparative studies Leguminosae (Baretta-Kuipers, 1981; Herendeen, 2000; Gasson et al., 2003), and especially with the early diverging Detarioideae In order to identify material as precisely as possible, initially, we (Cowan, 1981; Polhill, 1994; Bruneau et al., 2014). The three fossil searched the Inside Wood Database (IWD; http://insidewood.lib.ncsu. wood specimens may be subdivided into two types, differing most edu/), an online database containing 9072 modern and fossil wood de- significantly in terms of the size of the axial canals. As elaborated scriptions (last accessed 5 June 2017), using different combinations of below, comparison with collections at RBGK allows assignment to features that we considered to be of diagnostic value. In these searches the Prioria-clade sensu Breteler (1999), an emended supergenus

Plate I. Prioria hodgesii sp. nov., Rodríguez-Reyes, Gasson, Falcon-Lang et Collinson. STRI 14165 (holotype).

1. Growth rings indistinct, wood diffuse porous. STRI 14165. TS, scale: 1 mm. 2. Vessels solitary and in radial multiples of 2–3. Parenchyma in irregularly spaced bands (arrows). STRI 14165, TS, scale: 500 μm. 3. Simple perforation plates (SPP). STRI 14165, RLS, scale: 100 μm. 4. Intervessel pitting alternate (IVP). STRI 14165, TLS, scale: 100 μm. 5. Intervessel pitting polygonal. STRI 14165, TLS, scale: 25 μm. 6. Vessel–ray pitting (VRP) similar to intervessel pits (IVP). STRI 14165, RLS, scale: 50 μm. 7. Detail of vessel–ray pitting. STRI 14165, RLS, scale: 100 μm. 8. Thin-to thick-walled fibres. STRI 14165, TLS, scale: 100 μm. 9. Rays 1–3-seriate and non-septate fibres (NSF). STRI 14165, scale: 100 μm. 10. Irregular parenchyma bands 3–6 cells wide. Occasional paratracheal aliform parenchyma (arrow). STRI 14165, TS, scale: 100 μm.

Plate II. Prioria hodgesii sp. nov., Rodríguez-Reyes, Gasson, Falcon-Lang et Collinson. STRI 14165 (holotype). (see on page 6)

1. Axial parenchyma strands N8 cells high (arrow). STRI 14165, TLS, scale: 100 μm. 2. Axial parenchyma strands 4 cells high (between arrows). STRI 14165, TLS, scale: 100 μm. 3. Detail of short axial parenchyma strands. STRI 14165, TLS, scale: 100 μm. 4. Rays 1–3-seriate with long tails seen in tangential section (arrows). STRI 14165, TLS, scale: 100 μm. 5. Axial paratracheal parenchyma aliform (arrows). STRI 14165, TS, scale: 100 μm. 6. Ray composed of procumbent cells (PC) and N4 marginal rows of square cells (SC). STRI 14165, RLS, scale: 100 μm. 7. Normal axial canals diffuse and in pairs (arrows). STRI 14165. TS, scale: 100 μm. 8. Rhomboidal crystals (arrows) in chambered axial parenchyma cells. STRI 14165, TLS, scale: 100 μm. 9. Crystals in chambered axial parenchyma cells (arrows). STRI 14165, TLS, scale: 100 μm. 10. Square ray cells with more than one crystal of about the same size per chamber (arrow). STRI 14165, RLS, scale: 100 μm.

Plate III. Prioria canalensis sp. nov., Rodríguez-Reyes, Gasson, Falcon-Lang et Collinson. STRI 36273 (holotype). (see on page 7)

1. Growth rings indistinct, marked by narrow bands of marginal parenchyma (arrows) and wood diffuse porous. TS, scale: 1 mm. 2. Vessels in radial multiples of commonly 2 –5. STRI 36273, TS, scale: 250 μm. 3. Perforation plates simple (arrow). STRI 36273, RLS, scale: 100 μm. 4. Intervessel pitting alternate. STRI 36273, TLS, scale: 100 μm. 5. Intervessel pitting minute to small. STRI 36273, TLS, scale: 50 μm. 6. Vessel–ray pitting (circle) similar to intervessel pits in size and shape. STRI 36273, RLS, scale: 100 μm. 7. Tyloses abundant (arrow). STRI 36273, TS, scale: 100 μm. 8. Tangential view of tyloses (arrow). STRI 36273, TLS, scale: 100 μm.

Plate IV. Prioria canalensis sp. nov., Rodríguez-Reyes, Gasson, Falcon-Lang et Collinson. STRI 36273 (holotype). (see on page 8)

1. Non-septate fibres (NSF). STRI 36273, TLS, scale: 100 μm. 2. Minutely bordered pits (arrows) common in radial walls of fibres. STRI 36273, RLS, scale: 100 μm. 3. Fibres thin to thick walled and vessels with vasicentric winged-aliform parenchyma (arrows). STRI 36273, TS, scale: 100 μm. 4. Marginal band of parenchyma 1–4 cells wide (arrows). STRI 36273, TS, scale: 100 μm. 5. Parenchyma strands 6 cells high and abundant crystals present in chambered cells. STRI 36273, TLS, scale: 50 μm. 6. Rays typically 3-seriate and fibres pitted in tangential walls. STRI 36273, TLS, scale: 100 μm. 7. Ray body composed of procumbent cells (PC) and 1 row of square cells (SC). STRI 36273, RLS, scale: 100 μm. 8. Solitary rhomboidal crystal in square ray cell (arrow). STRI 36273, RLS, scale: 50 μm. 9. Diffuse normal axial canals (arrows). STRI 36273, TS, scale: 100 μm. 10. Short line of normal axial canals (arrow). STRI 36273, TS, scale: 200 μm. 48 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61

Plate I. O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 49

Plate II. (captiononpage4) 50 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61

Plate III. (captiononpage4) O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 51

Plate IV. (captiononpage4) 52 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 that absorbs Prioria sensu stricto, Kingiodendron Harms, Oxystigma b1 mm high; normal axial canals diffuse, in short tangential lines and Harms and Gossweilerodendron Harms.Weemphasisethatitisthe occasionally, in long tangential lines with a mean tangential diameter emended supergenus Prioria,ratherthanPrioria Grisebach, 1860 of 182 μm; rhomboidal solitary crystals abundant in chambered paren- sensu stricto, to which we refer our fossils. Fossil wood of this chyma cells. supergenus has not been, hitherto, described, and because the new Holotype: STRI 36273, designated herein, consisting of a hand speci- material differs from all extant Prioria-clade wood (Banks and men and a total of three thin sections. The original trunk in the field had Gasson, 2000), it is referred to two new species as follows: a preserved length of 2 m and 0.8 m diameter. Prioria hodgesii Rodriguez-Reyes, Gasson, Falcon-Lang et Collinson, Other material: STRI 36400, consisting of a hand specimen and thin sp. nov. (Plates I–II) sections. Diagnosis: Growth rings indistinct, wood diffuse-porous; perfo- Etymology: specific epithet canalensis refers to the Panama Canal, ration plates simple; intervessel pitting alternate; non-septate fi- where the fossil woods were collected. bres; axial parenchyma in discontinuous tangential bands 3–6 Description: Growth rings marked by narrow bands of marginal cells wide; paratracheal axial parenchyma, locally aliform; rays 1– parenchyma; wood diffuse-porous (Plate III, 1); vessels solitary 3-seriate; normal axial canals mostly diffuse with occasional short (57%) or in radial multiples of 2–5(−7) (Plate III, 2); vessel outline tangential lines and a mean diameter of 86 μm; abundant crystals oval; perforation plates simple (Plate III, 3); intervessel pits alternate in chambered cells in the axial parenchyma strands. (Plate III 4, 5) and minute to small (mean pit diameter 5.0 ± 0.99 μm, Holotype: STRI 14165, designated here, comprising a hand specimen n = 25; range 3–7 μm) (Plate III,5);vessel–ray pits with distinct bor- and three sections in TS, TLS and RLS. The preserved diameter of the axis ders, similar in diameter (mean 4.0 ± 1.5 μm, n = 25; range 2–7 μm) is 0.12 m. and shape to intervessel pits (Plate III, 6) (mean tangential vessel di- Repository: Center for Tropical Paleoecology and Archaeology, ameter 137 ± 23 μm (n = 50; range 90–190 μm); mean vessel den- Smithsonian Tropical Research Institute, Panama. sity 9.7 ± 3.1 per mm2 (n = 15; range 6–18 per mm2); mean vessel Type locality: Hodges Hill (Gaillard Cut of Panama Canal) near element length 308 ± 72 μm (n = 25; range 210–540 μm); tyloses Paraiso, Panama City, Panama (Latitude 09°02′51.75″N; Longitude occasionally present (Plate III,7,8));vasculartracheidsnot 79°39′14.02″W). observed. Stratigraphic horizon: ~20 m above the base of the Cucaracha Forma- Fibres, non-septate (Plate IV, 1) with minutely bordered pits com- tion (Gaillard Group); Lower Miocene (19–18.5 Ma). mon on radial walls and occasional in tangential walls (Plate IV,2);fi- Etymology: the specific epithet refers to the type locality of Hodges bres, thin- to thick-walled (Plate IV,3). Hill. Paratracheal axial parenchyma, vasicentric to short winged- Description: Growth rings indistinct; vessels diffuse-porous, 49% sol- aliform (Plate IV); occasional marginal bands of parenchyma 1–4 itary and 51% in radial multiples of 2–3(−5) (Plate I,1,2);vesselswith cells wide (Plate IV, 4); axial parenchyma strands 3–8 cells high oval outline (Plate I, 2); perforation plates, simple (Plate I,3); (Plate IV,5). intervessel pits alternate, polygonal and of small to medium size Rays, heterocellular, 1–3(−4)-seriate (mean 2.8 ± 0.7, n = 50) (mean 7.0 ± 1.1, n = 25; range 4–8 μm) (Plate I, 4, 5); vessel–ray pitting (Plate IV, 6) and commonly b1 mm (mean 0.8 ± 0.2 mm, n = 50; similar in diameter and shape to intervessel pits (mean 5.0 ± 1.4 μm, n range 0.4–1.5 mm; mean ray spacing 9.1 ± 2.5 per mm, n = 15; =25;range2(–8) Plate I, 6, 7); mean tangential vessel diameter 135 ± range 5–13 per mm). Rays either composed of procumbent bodies and 24.6 μm (n = 50; range 90–180 μm); mean vessel density 12 ± 4.5 per one or two rows of square cells (Plate IV, 7) or occasionally composed mm2 (n = 10; range 6–18) (Plate I, 1); mean vessel element length 520 exclusively of procumbent cells. ± 177.7 μm (n = 25; range 220–840 μm); tyloses absent; vascular tra- Solitary rhomboidal crystals found in square ray cells (Plate IV, 8), cheids not observed. but mostly abundant in chambered parenchyma strands (Plate IV, 5). Fibres, thin- to thick-walled (Plate I,8),non-septate(Plate I, 9); pits There are also occasional rhomboidal crystals in non-chambered paren- difficult to observe, but with, probably, simple to minutely bordered pits chyma strands. common on radial and tangential walls. Intercellular normal axial canals diffuse (Plate IV, 9) and commonly Axial parenchyma, mostly in irregularly spaced bands of 3–6 cells forming short and, rarely long tangential lines (Plate IV, 10) that alter- wide (Plate I, 10); occasional diffuse apotracheal and aliform nate with the marginal parenchyma bands. Axial canals mean diameter paratracheal parenchyma present (Plate I,10;Plate II, 5). Axial paren- 182 ± 32 μm (n = 30; range 111–242 μm). chyma strands with a highly variable number of cells (Plate II,1,2,3), IAWA features numbers present: 2, 5, 13, 22, 24v, 25, 30, 42, 47, 52, but mostly over 8-cells high (Plate II,1). 56v, 61, 66, 69, 79, 80, 82, 89, 92 V, 93, 94, 97, 98v, 104v, 106, 107, Rays, heterocellular, 1–3seriate(Plate II,1,2,3,4),rarelyupto4 127v, 128, 129v, 137, 141, 142. cells wide (Plate II,1)andb1 mm high (mean height 0.5 ± 0.2 mm, n = 50; range 0.25–0.92); rays commonly composed of procum- 5. Identification of fossil wood types bent cells and 2–4 rows of upright and/or square marginal cells (Plate II, 6). Mean ray spacing 11 ± 2 per mm (n = 15; range 7–14). In order to identify the woods, initially, we searched the Inside Solitary rhomboidal prismatic crystals, abundantly present in cham- Wood Database (IWD; http://insidewood.lib.ncsu.edu/) and then criti- bered axial parenchyma cells, forming chains (Plate II, 8, 9). Solitary cally assessed the likelihood of each potential match by comparing the crystals also present in square ray cells (Plate II, 10). fossil woods with images of modern woods in the IWD as well as Intercellular normal axial canals diffuse (Plate II,7)and,locally, through comparison with material in the Jodrell Laboratory of the forming short tangential lines (Plate II, 7); mean axial canal diameter Royal Botanic Gardens, Kew, U.K. (RBGK). 86 ± 20 μm (n = 30; range 51–135). IAWA feature numbers: 2, 5, 13, 22, 23, 26, 30, 42, 47, 53, 61, 66, 69v, 5.1. IWD search 1: affinities with Leguminosae 76v, 85, 91, 92, 93, 94v, 97, 106, 107v, 141v, 142, 154. Prioria canalensis Rodriguez-Reyes, Gasson, Falcon-Lang et In the first search, we input only one relatively unusual character ob- Collinson, sp. nov. (Plates III–IV) served in both wood types: normal axial canals in short tangential lines Diagnosis: Growth rings marked by narrow bands of marginal paren- (128 p). Most results were spread through the Dipterocarpaceae chyma; wood diffuse-porous; perforation plates simple; intervessel (Anisoptera, Dipterocarpus, Dryobalanus, Hopea, Shorea and Parashorea), pitting alternate and minute to small; axial parenchyma winged-aliform Lauraceae (Cryptocarya), Rosaceae (Prunus) and Vochysiaceae (Vochysia), or in narrow bands up to 3 cells wide; rays 1–3(−4)-seriate and with the remainder within Leguminosae. The Dipterocarpaceae matches O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 53 could be excluded because they show, mostly, solitary vessels, very wide 2003), however, this is a highly problematic and uncertain feature, es- rays (4–10-seriate) and abundant aliform paratracheal parenchyma, pecially if the intervessel pits are minute. In fossil woods, it is difficult characters absent in both fossil wood types. The single Lauraceae repre- to unambiguously determine the presence of vestured pits, due to min- sentative, Cryptocarya, can also be ruled out because it possesses large eral deposits that can resemble or obscure any vesturing (Wheeler and intervessel pits and prominent oil cells associated with axial parenchyma, Baas, 1992). features not observed in the fossils. Of the other matches, Rosaceae (Pru- In terms of the new phylogeny (LPWG, 2017), the records of the initial nus) has distinct growth rings, vessels in diagonal pattern, helical thicken- IWD search include 12 genera (, , Brandzeia, Copaifera, ings and intercellular canals of traumatic origin while Vochysiaceae , , , Kingiodendron, Librevillea, Prioria, Sindoropsis, (Vochysia) commonly has large vessels, abundant paratracheal parenchy- ) that all fall within the early-diverging Detarioideae subfami- ma and intercellular canals of traumatic origin, both significantly different ly. The Detarioideae comprises 84 genera and is pantropical in distribu- from the fossil wood types. tion (Lewis et al., 2005; LPWG, 2017). In a survey of the IWD and key However, the fossils do show a close match with the remaining le- literature (Wheeler and Baas, 1992; Herendeen, 2000; Gasson et al., gume taxa in the search. In the current version of IWD (accessed 5 June 2003), we confirmed this association, finding the common occurrence 2017), these legume records remain coded in terms of the traditional le- of the key characters in the fossil woods and members of the subfamily gume phylogeny, and assigned to the former caesalpinioid-grade. This Detarioideae (e.g., aliform parenchyma combined with marginal narrow former caesalpinioid-grade has a lot of variation in its wood anatomy, bands, thin to thick walled fibres, narrow non-storied rays, prismatic crys- probably because this group is paraphyletic (see Gasson et al., 2003,for tals abundant in chambered parenchyma cells). Hence we focused our an extensive and detailed study of this grouping), but a few traits com- comparison on this subfamily. monly occurring in them are: winged-aliform paratracheal parenchyma, confluent and marginally banded parenchyma, medium to thick walled 5.2. IWD search 2: affinities with subfamily Detarioideae fibres, non-storied biseriate rays (homocellular or with a row of square or upright cells), vestured pits and chains of crystals in chambered paren- In order to identify the fossil woods more precisely, we conducted a chyma (Metcalfe and Chalk, 1950; Wheeler and Baas, 1992; Gasson, second IWD search using a set of ten features that occur in all three spec- 1994; Gasson et al., 2003; Wheeler and Manchester, 2002; Brandes and imens as follows: wood diffuse-porous (5 p), perforation plates simple Barros, 2008; Pujana et al., 2011). (13 p), intervessel pitting alternate (22 p), vessel–ray pitting with distinct The alternate intervessel pitting seen in the fossils is characteristic of borders and similar to intervessel pitting in size and shape (30 p), fibres all of the Leguminosae but, it has been noted as a feature of diagnostic non-septate (66 p), paratracheal parenchyma vasicentric (79 p), potential because some groups can be separated by size of intervessel apotrachealparenchymainmarginalbands(89p),rays1–3-seriate (97 pitting, especially in the former Caesalpinioideae. For instance, large p), normal axial canals in short tangential lines (128 p) and prismatic intervessel pits are found in the Caesalpinia, Cassinae and Amhertisia crystals in chambered axial parenchyma strands (142 p). groups. The presence of vestured pits is another key trait for this sub- The results of this search comprised eight genera (Anthonotha, family in different surveys (e.g., Herendeen et al., 2003; Gasson et al., Bikinia, Copaifera, Detarium, Eperua, Prioria, Sindoropsis, Tessmannia),

Table 1 Relevant wood anatomical properties (traumatic and normal axial canals, storied structure of rays) for key representatives of the subfamily Detarioideae, and Prioria sensu Breteler (1999) in particular. (Data from Banks and Gasson, 2000; Inside Wood Database: http://insidewood.lib.ncsu.edu/.)

Genus Traumatic canals Normal axial canals (distribution) Storied structure of rays (type)

Tribe Detarieae Anthonotha P. Beauv. Present Present (short tangential lines) Absent Augouardia Pellegr. Absent Absent Absent Bikinia Present Present (short tangential lines) Absent Benth. Present Absent Present (irregular) Brandzeia Braill. Present Present (marginal tangential lines) Absent Colophospermum Kirk ex Benth. Absent Absent Absent Copaifera Lindl. Absent Present (long tangential lines) Absent Benth & Hook. Absent Absent Present Daniellia Benn. Absent Present (diffuse) Present Detarium Juss. Absent Present (long tangential lines) Absent Eperua Aubl. Present Present (long tangential lines) Absent Benn. Present Absent Absent Roxb. Absent Absent Absent Pellegr. Present Absent Present (irregular) Librevillea Hoyle Present Present (diffuse) Absent Hauman Present Present (diffuse) Absent A. Chev. Present Absent Absent Neoapaloxylon S. Rauschert Absent Absent Absent Oddoniodendron DeWilld. Present Present (diffuse) Absent Vogel Present Absent Present (irregular) Pseudosindora Dewit. Absent Present (long tangential lines) Absent Sindora Miq. Absent Present (long tangential lines) Absent Sindoropsis J. Leonard Absent Present (long tangential lines) Absent Stemoabsentcoleus Harms Absent Absent Absent Tessmannia De Wild. Absent Present (short, long tangential lines) Absent Hauman Present Absent Absent

Prioria emended Breteler 1999 Gossweilerodendron Harms Present Present (diffuse, short tangential lines) Absent Kingiodendron Harms Present Present (diffuse, short tangential lines) Absent Oxystigma Harms Present Present (diffuse) Absent Prioria Griseb. Present Present (diffuse, short tangential lines) Absent 54 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61

Table 2 Comparison of major characteristics for the fossil woods and the most similar modern woods.

Taxon Prioria hodgesii Kingiodendron alternifolium Prioria canalensis

Specimen STRI 14165 RBGK Philippines 1954 STRI 36273 RBGK IFI 3412 Growth rings Indistinct Indistinct Variable; marked by narrow bands of Variable; marked by bands of marginal marginal parenchyma parenchyma Vessel Diffuse-porous Diffuse-porous Diffuse-porous Diffuse-porous porosity Vessel 2–3(−5); 49% solitary 2–5; 49% solitary 2–5(−7); 57% solitary 2–5; 61% solitary arrangement and grouping Perforation Simple Simple Simple Simple plates Intervessel pit Alternate Alternate, polygonal Alternate Alternate; polygonal? arrangement Intervessel pit 7 ± 1.1 (range 4–8) 1–6 5 ± 0.99 (range 3–7) 3–8 diameter (μm) Vessel–ray pit similar to ivp similar to ivp similar to ivp similar to ivp shape Vessel–ray pit 5 ± 1.4 (range 2–8) 4 ± 1.3 (range 2–7) 4 ± 1.5 (range 2–7) 4 ± 1.4 (range 2–6) diameter (μm) Mean 135 ± 25 (range 90–180) 199 ± 29 (range 121–245) 137 ± 23 (range 90–190) 162 ± 22 (range 120–190) tangential vessel diameter (μm) Vessels per 12 ± 4.5 (range 6–18) 4.5 ± 1.1 (range 3–6) 9.7 ± 3.1 (range 6–18) 3.9 ± 1.0 (range 2–5) mm2 Mean vessel 520 ± 178 (range 220–840) 364 ± 76 (range 230–483) 308 ± 72 (range 210–540) 429 ± 92 (range 244–644) element length (μm) Fibre pits Simple to minutely bordered; Simple to minutely bordered; Simple to minutely bordered; common only Simple to minutely bordered; common only common on radial and tangential common only on radial walls on radial walls on radial walls walls Septate fibres Absent Absent Absent Absent Fibre wall Thin- to thick-walled Thin- to thick-walled Thin- to thick-walled Thin- to thick-walled thickness Paratracheal Occasionally aliform Vasicentric, occasionally aliform Vasicentric to short winged-aliform Scanty, vasicentric, short winged-aliform and axial confluent parenchyma Apotracheal Occasionally diffuse Absent Absent Absent axial parenchyma Banded axial Bands irregularly spaced 3–6 Bands more than 3–cells wide Occasional marginal bands 1–4 cells wide Narrow bands of 3–4 cells wide parenchyma cells wide Axial Variable, mostly N8 cells 2–4 cells 3–8 cells 2–4 cells parenchyma strand length Ray width 1–3(−4) cells 1–3 cells 1–3(−4) cells 1–3 cells (cells) Ray height 0.5 ± 0.2 (range 0.3–0.9) no data 0.8 ± 0.2 (range 0.4–1.5) 0.5 ± 0.3 (range 0.2–1.5) (mm) Ray cellular Commonly with procumbent Body ray cells procumbent with one Procumbent bodies and one or two rows of Procumbent bodies with 2–4 rows of composition bodies and 2–4 rows of upright row of square and upright cells; square ray cells; occasionally composed of upright cells and/or square marginal cells occasionally 2–4rowsalsopresent exclusively procumbent cells Sheath cells Absent Absent Absent Absent Tile cells Absent Absent Absent Absent Perforated ray Absent Absent Absent Absent cells Rays per linear 11 ± 2 (range 7–14) 8.6 ± 1.3 (range 6–10) 9.1 ± 2.5 (range 5–13) 6 ± 0.9 (range 4–7) millimetre Storied Absent Absent Absent Parenchyma strands irregularly storied structures Intercellular Diffuse and in short tangential Diffuse; occasionally in short Diffuse and commonly forming short Axial canals diffuse and occasionally forming normal axial lines tangential lines tangential lines; occasional long lines that short and long tangential lines that alternate canals alternate with the marginal parenchyma with the marginal parenchyma bands. bands Intercellular 86 ± 20 (range 51–135). 172 ± 52 (range 85–242) 182 ± 32 (range 111–242). 183 ± 33 (range 142–245) axial canal diameter (μm) Prismatic Present in chambered axial Abundant in chambered axial Abundant in chambered axial parenchyma Present in chambered axial parenchyma cells crystals parenchyma cells and forming parenchyma cells cells, but also present in square ray cells and in square ray cells chains O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 55 all within the Detarioideae subfamily. In order to discriminate between 5.3. Comparison with Prioria emended Breteler (1999) Detarioideae genera, and to find the best match for the fossils analysed here, we initially focused on four features: The genus Prioria underwent significant systematic revision, with spe- cies formerly placed in Oxystigma, Kingiodendron, Gossweilerodendron, (i) Paratracheal parenchyma. The parenchyma is predominantly and Prioria, being incorporated within the emended genus, Prioria, paratracheal in most legumes (Gasson et al., 2003; Pujana et al., following the work of Breteler (1999). Oxystigma, Kingiodendron, and 2011). In all of the groups in Detarioideae the parenchyma Gossweilerodendron are endemic to equatorial Africa whereas Prioria tends to be a combination of vasicentric and winged-aliform, sensu stricto is found only in tropical Central America, so this revision is but usually combined with abundant broad (more commonly not in accord with biogeographic distribution, and suggests an early fi in the , Detarium and groups) and narrow diversi cation (De la Estrella et al., 2017). It is the emended genus Prioria bands of parenchyma (commonly in the , (Breteler, 1999), rather than Prioria Grisebach, 1860, to which we refer and Group). our fossils. The occurrence of normal axial canals diffusely arranged and (ii) Storied structures. The occurrence of storying in rays, parenchy- in short tangential lines is restricted to the emended Prioria (Gasson et fi ma and fibres is highly variable in the Leguminosae, although al., 2003), which fully justi es the assignment of the Panama Canal ‘ ’ ‘ ’ this is a feature of great diagnostic potential. In Detarioideae, sto- woods to the group containing Prioria sensu stricto, Gossweilerodendron , ‘ ’ ‘ ’ ried rays are rare, but in some genera, such as Daniellia, all the Kingiodendron and Oxystigma . rays are storied. Although the three fossil wood specimens share several key features, (iii) Prismatic crystals. Prismatic crystals are dominant in apotracheal the size and distribution of normal axial canals suggest that two different parenchyma cells in most of the members of the Detarioideae. species are involved. Prioria hodgesii sp. nov. possesses normal axial canals μ There are some exceptions, for example, crystals are rarely ob- with a mean diameter of 86 m, which are mostly diffuse with occasional ‘ ’ served in , Copaifera and Ecuadendron (Gasson et al., short tangential lines, a pattern typical for Kingiodendron but also seen in ‘ ’ ‘ ’ 2003) or they are observed mainly in the ray cells (Plagiosiphon, Prioria sensu stricto and Gossweilerodendron (Table 1). In contrast, Neochevalierodendron). Prioria canalensis sp. nov. possesses larger normal axial canals with a μ ‘ ’ (iv) Normal axial canals. A key feature in the fossils is the occurrence mean diameter of 182 m, which is most commonly seen in Prioria of normal axial canals distributed diffusely and in short tangen- sensu stricto. Prioria canalensis has diffuse, short and occasional long tial lines. Gasson et al. (2003) and Baretta-Kuipers (1981) state lines of normal axial canals. A detailed quantitative comparison between that only some genera of Detarioideae possess axial canals in var- the fossil taxa and the most closely similar extant material in the RBGK iable distribution patterns (Table 1). collection is presented in Table 2, and descriptive and illustrated compar- ison is given as follows: Prioria hodgesii sp.nov.sharesmostcharacterswithsomeRBGKspec- Reviewing the IWD search short list, Copaifera, Detarium, Eperua, imens of ‘Kingiodendron’ such as: growth rings indistinct, wood diffuse and Sindoropsis show normal axial canals in long tangential lines porous (Plate V, 1); perforation plates simple (Plate V, 2); intervessel only, different from the fossils. Tessmannia has normal axial canals pitting alternate ( Plate V, 3) and mostly small-sized (Plate V, 4); vessel– arranged in long and short lines, a combination not seen in the fos- ray pitting similar in diameter and shape to intervessel pits (Plate V,5); sils. Anthonotha and Bikinia do show normal axial canals in short tan- fibres thin-to-thick walled (Plate V, 6); non-septate fibres (Plate V,7); gential lines but not with the diffuse distribution seen in the fossils. rays 1–3-seriate (Plate V, 7); parenchyma in discontinuous tangential Of the short-listed taxa, the fossils share most features with Prioria, bands 3–6 cells wide (Plate V; 1, 8) abundant crystals in chambered and therefore a detailed comparison with this genus was undertaken cells in the axial parenchyma strands (Plate V,2);diffusenormalaxialca- (Table 2). nals present (Plate V, 1) and occasionally, in short lines (Plate V,8).

Plate V. Comparative material of ‘Kingiodendron’ alternifolium. RBGK Philippines 1954.

1. Growth rings indistinct, wood diffuse porous. Normal axial canals diffuse (arrows). RBGK Philippines 1954, TS, scale: 100 μm. 2. Simple perforation plates (spp). Abundant rhomboidal crystals (arrow). RBGK Philippines 1954, RLS, scale: 100 μm. 3. Intervessel pitting (ivp) alternate. RBGK Philippines 1954, TLS, scale: 50 μm. 4. Intervessel pitting minute. RBGK Philippines 1954, TLS, scale: 10 μm. 5. Vessel–ray pitting (vrp) similar to the intervessel pitting in size and shape. RBGK Philippines 1954, RLS, scale: 25 μm. 6. Fibres thin-to-thick walled. RBGK Philippines 1954, TS, scale: 25 μm. 7. Rays 1–4-seriate. Fibres non septate. RBGK Philippines 1954, TLS, scale: 100 μm. 8. Normal axial canals in pairs (arrow). RBGK Philippines 1954, TS, scale: 100 μm.

Plate VI. Comparative material of ‘Prioria’ copaifera. RBGK IFI 3412, RBGK W-16558. (see on page 14)

1. Growth rings marked by marginal parenchyma bands (arrows). RBGK IFI3412s, scale: 100 μm. 2. Vessels in radial multiples of commonly 2–6 and axial canals in pairs (arrows). RBGK IFI 3412, scale: 100 μm. 3. Perforation plate simple (arrow). RBGK W-16558, scale: 100 μm. 4. Intervessel pitting alternate. RBGK IFI 3412, scale: 20 μm. 5. Vessel–ray pitting similar to intervessel pits. RBGK IFI 3412, scale: 40 μm. 6. Non-septate fibres. Arrows mark fibre pits in tangential walls. RBGK IFI 3412, scale: 100 μm. 7. Fibres with simple to minutely bordered pits common in radial walls. RBGK W-16558, scale: 100 μm.

Plate VII. Comparative material of Prioria copaifera. RBGK IFI 3412, RBGK W-16558. (see on page 15)

1. Axial parenchyma ranging from vasicentric to winged-aliform (arrows). RBGK IFI3412, scale: 100 μm. 2. Rays 1–2 (3)-seriate. RBGK IFI 3412, scale: 100 μm. 3. Rays composed of procumbent cells (PC) and 1 row of marginal square cells (SC). RBGKW-16558, scale: 100 μm. 4. Solitary rhomboidal crystals (arrows) occasional in square ray cells. RBGK IFI 3412, scale: 100 μm. 5. Rhomboidal crystals abundant (arrows) in parenchyma chambered cells. RBGK IFI 3412, scale: 50 μm. 6. Diffuse normal axial canals (arrows). RBGK IFI 3412, scale: 50 μm. 7. Short line (2) of normal axial canals. RBGK IFI 3412, scale: 50 μm. 56 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61

Plate V. O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 57

Plate VI. (captiononpage12) 58 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61

Plate VII. (captiononpage12) O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 59

Prioria canalensis sp. nov. shares most characters with some RBGK focus the discussion at generic level, organized by geological age specimens of ‘Prioria’ as follows: Growth rings marked by marginal from oldest to youngest, as follows: bands of parenchyma (Plate VI,1,2);wooddiffuse-porous(Plate VI,1); fi perforation plates simple (Plate VI, 3); intervessel pits alternate and min- (i) The oldest wood of secure Detarioideae af nity is reported from ute to small (Plate VI, 4); vessel–ray pits similar in diameter and shape to the Eocene of France, Aulacoxylon (De Franceschi and De Ploëg, intervessel pits (Plate VI,5);fibres non-septate (Plate VI,6)withminute- 2003); it has normal axial canals, but all the rays are storied, so ly bordered pits common in radial walls and occasional in tangential it differs from the new fossils. walls (Plate VI, 6, 7); paratracheal axial parenchyma vasicentric to short (ii) Oligocene reports of Detarioideae wood are much more common winged-aliform (Plate VII, 1); Rays heterocellular 1–2 (3) seriate (Plate and include Erythrophloeoxylon (Boureau, 1957; Müller-Stoll and VII, 2), solitary rhomboidal crystals found in square ray cells (Plate VII, Mädel, 1967), Cynometroxylum (Delteil-Desneux, 1981), 3, 4), but mostly abundant in chambered parenchyma strands (Plate Afzelioxylon (Louvet, 1965), Copaiferoxylon (Fessler-Vrolant, fi VII, 5). There are also occasional rhomboidal crystals in non-chambered 1977), and Detarioxylon (Boureau and Louvet, 1970). The rst parenchyma strands. Intercellular normal axial canals diffuse (Plate three taxa do not possess normal axial canals. Detarioxylon fl VII, 6) and forming short lines (Plate VI,2;VII, 7) that alternate with has normal axial canals, and also distinct con uent parenchy- – the marginal parenchyma bands (Plate VI,1). ma and rays 4 10-seriate. Copaiferoxylon also has normal axial We also note some differences between these extant taxa and canals, but these are mostly distributed in long tangential thefossilssuchasthemorecommonconfluent axial parenchyma lines, and it has winged-aliform parenchyma. As such, all this and marginal upright cells composing the rays in the modern Oligocene material differs from the new fossil material report- wood contrasting with the rarely present confluent parenchyma ed here. and square marginal cells in the rays. In addition, in the extant spe- (iii) Miocene reports of Leguminosae fossil woods are numerous cies, the diffuse normal axial canals are considerably wider than the (Wheeler and Baas, 1992). They include Acrocarpus (Yadav, vessels, a feature that is not seen in the fossils (Table 2). 1989), Acrocarpoxylon (Gottwald, 1994), Afzelioxylon (Lemoigne et al., 1974; Lemoigne, 1978), Brachystegioxylon 6. Comparison with previously described fossil woods (Lakhanpal and Prakash, 1970), Gleditsia (Prakash et al., 1962; Watari, 1952), Peltophoroxylon (Lemoigne, 1978; In order to demonstrate, more thoroughly, the novelty of the new Ramanujam, 1954, 1960), and Koompassioxylon (Bande and fossil species, we compare them with (i) fossil woods previously de- Prakash, 1980; Yadav, 1989; Awasthi and Mehrotra, 1990). fi scribed from the Miocene Cucaracha Formation of Panama and with Miocene woods speci cally related to Detarieae include (ii) previously described fossil woods with an affinity with Leguminosae Crudioxylon (Pons, 1980), Cynometroxylon (Chowdhury and subfamily, Detarioideae. Ghosh, 1946; Navale, 1959; Prakash and Awasthi, 1971; Trivedi and Ahuja, 1978; Prakash, 1979; Bande and Prakash, 6.1. Fossil woods from the Cucaracha Formation 1980; Vozenin-Serra et al., 1989; Delteil-Desneux, 1981; Sayadi, 1973), Kingiodendron (Awasthi and Prakash, 1987), A large collection of fossil wood has been studied recently from the Pahudioxylon (Chowdhury et al., 1960; Navale, 1962; Cucaracha Formation; however, no other specimens resemble the new Prakash and Awasthi, 1971; Prakash and Tripathi, 1975; material reported here (Rodríguez-Reyes, 2014). In an earlier phase of Louvet, 1974; Lorch and Fahn, 1959; Müller-Stoll and Mädel, investigation, Berry (1918) collected large trunks from the Cucaracha 1967), and Sindoroxylon (Lemoigne et al., 1974). Most of Formation (and the upper part of the underlying Culebra Formation) these fossil woods lack normal axial canals or at least they during the original construction of the Panama Canal. Berry (1918) are not mentioned in the literature, consequently, we rule identified a few of these specimens as Taenioxylon multiradiatum. them out as matches for the new fossils. Sindoroxylon does Taenioxylon is a palaeotaxon established and reported by Felix (1882, possess normal axial canals, but it tends to have more abun- fl 1883, 1886, 1887) from several localities in Central America, Asia and dant con uent parenchyma, frequently forming bands over 3 Europe. The resolution of the images in the publication is somewhat cells wide. Various other Miocene woods of Detarioideae are poor, but adequately shows that there are no axial canals and nor are assigned to : Bauhinia deomalica (Awasthi and there abundant crystals in these specimens. Neither are these features Prakash, 1987; Agarwal, 1991; Awasthi, 1992), Bauhinia mentioned in the description. Although Berry emphasises the poor tertiaria (Awasthi and Mehrotra, 1989), Bauhinia indicum preservation of most of the specimens, and the limitations of the pre- (Kramer, 1974a), and Bauhinia miocenicum (Trivedi and pared sections, we consider these woods to be different from our Prioria Panjwani, 1986). All of these woods lack axial canals and specimens from Hodges Hill, at least, based on the observation of the most also show storied rays and axial parenchyma strands. only illustration available in Berry's publication. Also Taenioxylon has Many of these woods are also reported from the Pliocene. later been recognized as synonym of Pahudioxylon (Chowdhury et al., (iv) Pliocene reports include Caesalpinioxylon (Koeniguer, 1973, fi 1960; Navale, 1962; Prakash and Awasthi, 1971; Prakash and Tripathi, 1974), which has septate bres, banded parenchyma (more 1975; Louvet, 1974; Lorch and Fahn, 1959; Müller-Stoll and Mädel, than 3 cells wide) and exclusively uniseriate and storied 1967), a different Leguminosae taxon. rays. There are also reports of Saracoxylon (Du, 1988)and Cynometroxylon (India: Lakhanpal et al., 1984; Guleria, 1984; 6.2. Fossil woods of Detarioideae Thailand: Vozenin-Serra and Privé-Gill, 1989), which are probably Detarieae but lack the normal axial canals seen in The fossil record of Leguminosae woods is more extensive than our new fossils. for any other eudicot family. Unfortunately many fossil woods assigned to this family cannot be certainly placed into any particular This comprehensive review of Detarioideae fossil woods demon- extant group or genus and are named Leguminoxylon.Forcompre- strates the uniqueness of the new species of Prioria described here. hensive and extensive surveys of the fossil wood record for Leguminosae, see the following reviews: Müller-Stoll and Mädel 7. Biogeographical and palaeoecological implications (1967), Crepet and Taylor (1985), Crepet and Herendeen (1992), Wheeler and Baas (1992),andHerendeen (2000).Hereweconcen- Prioria hodgesii sp.nov.andPrioria canalensis sp.nov.areclosely trate our comparison on woods in the Detarioideae subfamily. We similar woods, but distinct in terms of the size and distribution of 60 O. Rodríguez-Reyes et al. / Review of Palaeobotany and Palynology 246 (2017) 44–61 normal axial canals. Both morphotypes are similar to species in the Baretta-Kuipers, T., 1981. Wood anatomy of Leguminosae: its relevance to . In: Polhill, R.M., Raven, P.H. (Eds.), Advances in Legume Systematics, Part 2. Royal supergenus Prioria as emended by Breteler (1999), which were for- Botanic Gardens, Kew, pp. 677–805. merly placed in ‘Kingiodendron’ and ‘Prioria’ sensu stricto. These ex- Berry, E.W., 1918. The fossil higher plants from the Canal Zone. Bull. U.S. Nat. Mus. 15–44. tant genera are large trees adapted to seasonally-flooded riparian Boureau, E., 1957. Etude anatomique et dendroclimatologique d'un bois silicifiétertiaire des environs de Laruscade (Gironde), Leguminoxylon schoelleri. C. R. 82e Congr. Soc. environments. For example, Prioria copaifera Grisebach, up to Sav. Paris pp. 181–189. 40 m high, is found mostly in coastal areas of Nicaragua, Costa Boureau, E., Louvet, P., 1970. Sur deux espéces ligneuses tertiaries nouvelles de la région Rica, Panama and south to Colombia. The trees are often in nearly de Ouaou en Namous (Libye). Congr. Soc. Sav. Paris 3, 11–42. monospecific stands in coastal swamp forests and along estuaries Brandes, A.F.N., Barros, C.F., 2008. Wood anatomy of eight liana species of Leguminosae family from Atlantic Rain . Acta Bot. Bras. 22, 465–480. (Lewis et al., 2005), communities that are colloquially termed Brea, M., Franco, M.J., Lutz, A.I., 2012. Redescription and reassignment of Entrerrioxylon “cativales” (Giraldo Jimenez and del Valle Arango, 2011). Growing victoriensis from the Upper Miocene Paraná Formation, South America. Rev. – sites are often underwater during a portion of the year, along rivers Palaeobot. Palynol. 185, 13 25. Breteler, F.J., 1999. A revision of Prioria, including Gossweilerodendron, Kingiodendron, or in slopes lying inland form the coastal plain (Lopez and Kursar, Oxystigma and Pterygodium (Leguminosae-Caesalpinioideae-Detarieae) with empha- 1999; Grauel and Putz, 2004; Sandi and Flores, 2002). The typical sis on Africa. Wagening. Agric. Univ. Pap. 99, 1–61. distribution of this species is consistent with the geological context Bruneau, A., Klitgaard, B.B., Prenner, G., Fougère-Danezan, M., Tucker, S.C., 2014. Floral evolution in the Detarieae (Leguminosae): phylogenetic evidence for labile floral de- of the Hodges Hill locality, interpreted as a tidal estuarine envi- velopment in an early diverging legume lineage. Int. J. Plant Sci. 175, 392–417. ronment. Also, we have found other taxa related to those envi- Chowdhury, K., Ghosh, S., 1946. On the anatomy of Cynometroxylon indicum gen. et sp. nov., ronments, e.g., Malvaceae (Rodríguez-Reyes et al., 2014). a fossil dicotyledonous wood from Nailalung, Assam. Palaeobotanist 12, 435–447. Chowdhury, K.A., Ghosh, S., Kazmi, M., 1960. Pahudioxylon bankurensis gen. et sp. nov.: a Our observations provide compelling evidence for the presence of fossil wood from the Miocene bed of the Bankura District, West Bengal, India. Proc. the Detarioideae in Central America in Miocene times, and add to the Natl. Inst. Sci. India B26, 22–28. proposed wider distribution of the subfamily in the past (Brea et al., Cowan, R.S., 1981. Caesalpinioideae. In: Polhill, R.M., Raven, P.H. (Eds.), Advances in Le- gume Systematics, Part 1. Royal Botanic Gardens, Kew, pp. 57–64. 2012; Bruneau et al., 2014), extending the fossil record to Central Amer- Crepet, W., Herendeen, P.S., 1992. Papilionoid flowers from the early Eocene of southeast- ica and suggesting closer links with South American and African floras. ern North America. In: Herendeen, P.S., Dilcher, D.L. (Eds.), Advances in Legume Sys- We note that Detarioideae is much more diverse in Africa today (56% tematics, Part 4: The Fossil Record. Royal Botanic Gardens, Kew, pp. 43–55. fi fi of species) compared to the Neotropics, and we also note that Crepet, W.L., Taylor, D.W., 1985. The diversi cation of the Leguminosae: rst fossil evi- dence of the Mimosoideae and Papilionoideae. Science 228, 1087–1089. ‘Kingiodendron’, with which some of the features of our fossils are De Franceschi, D., De Ploëg, G., 2003. Origine de l'ambre des faciès sparnaciens (Éocène compared, is not distributed in the Neotropics today; its six species inférieur) du Bassin de Paris: le bois de l'arbre producteur. Geodiversitas 25, 633–647. are distributed through Indomalesia. These anomalies are consistent De la Estrella, M., Forest, F., Wieringa, J.J., Fougere-Danezan, M., Bruneau, A., 2017. Insights on the evolutionary origin of Detarioideae, a clade of ecologically dominant tropical with the emerging evolutionary and biogeographic understanding of African trees. New Phytol. 214, 1722–1735. Detarioideae, which implies an early origin (prior to the Eocene) and a Delteil-Desneux, F., 1981. Sur l'association de trois bois fossiles dans un même gisement more recent diversification event (earliest Miocene) that is slightly tertiaire du Djebel Nara (Tunisie centrale). Rev. Palaeobot. Palynol. 31, 289–310. Du, N., 1988. Fossil wood from the late Tertiary of Burma. Proc. K. Ned. Akad. Wet. A 91, older than the age of the fossils (De la Estrella et al., 2017). These molec- 213–236. ular biogeographic studies, together with our fossils, suggest that the Falcon-Lang, H.J., Wheeler, E.A., Baas, P., Herendeen, P.S., 2012. A diverse charcoalified as- current distribution of genera is relictual and does not faithfully reflect semblage of Cretaceous (Santonian) angiosperm woods from Upatoi Creek, Georgia, USA. Part 1: wood types with scalariform perforation plates. Rev. Palaeobot. Palynol. Miocene tropical biogeography. 184, 49–73. Farris, D.W., Cardona, A., Montes, C., Foster, D., Jaramillo, C., 2017. Magmatic evolution of Panama Canal volcanic rocks: a record of arc processes and tectonic change. PLoS One Acknowledgements 12, e0176010. Felix, J., 1882. Studien über fossile Hölzer. Inaugural Dissertation zur Erlangung der Doctorwurde. Universitatm Leipzig, Leipzig (84 pp.). ORR acknowledges funding from the Secretaría Nacional de Ciencia, Felix, J., 1883. Die fossilen Hölzer Westindiens. Samm. Palaeont. Abh. Teil 1 (1), 1–29. Tecnología e Innovación (SENACyT) (270-2010-108), Panama Canal Au- Felix, J., 1886. Untersuchungen über fossile Hölzer II. Z. Dtsch. Geol. Ges. 38, 483–492. Felix, J., 1887. Untersuchungen über fossile Hölzer III. Z. Dtsch. Geol. Ges. 39, 517–528. thority contract SAA-199520-KRP, U.S. National Science Foundation Fessler-Vrolant, C., 1977. Sur la presénce d'un nouveau bois fossile de légumineuse dans (NSF) Grants 0966884 (OISE, EAR, DRL) and 0824299 (EAR), National l'Oligocène de la Tunisie septentrionale. Actes 102e Congr. Nat. Soc. 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