The revised generic diagnosis, specific description and synonymy of the Late Cretaceous Rosannia manika from Alberta, Canada: Its phytogeography and affinity with family Lactoridaceae

Review of Palaeobotany and Palynology 159 (2010) 2–13

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Review of Palaeobotany and Palynology j o u r n a l h o m e p a g e : w w w. e l s e v i e r. c o m / l o c a t e / r e v p a l b o

The revised generic diagnosis, specific description and synonymy of the Late Cretaceous Rosannia manika from Alberta, Canada: Its phytogeography and affinity with family Lactoridaceae Satish K. Srivastava a,⁎, Dennis R. Braman b a b

Geology Consultant, 3054 Blandford Drive, Rowland Heights, California 91748-4825, United States Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta, Canada T0J 0Y0

a r t i c l e

i n f o

Article history: Received 18 March 2009 Received in revised form 13 October 2009 Accepted 15 October 2009 Available online 24 October 2009 Keywords: Rosannia manika Lactoripollenites pollen tetrads Late Cretaceous Lactoris fernandeziana Alberta, Canada

a b s t r a c t The genus Rosannia was erroneously diagnosed as a monad in Srivastava (1968a). It is re-diagnosed and its type species Rosannia manika is redescribed. The lost holotype is replaced by the surviving isotype as a lectotype here. Epitypes are designated and illustrated in this study to explain the morphology of R. manika in detail. Rosannia is an obligate ana-ulcerate tetrad with calymmate exine and granulose supratectal ornamentation. Its worldwide occurrence ranges from the Turonian to the Miocene. Lactoripollenites Zavada and Benson is a junior synonym of Rosannia. Pollen of extant Lactoris fernandeziana Phil. has a close morphological affinity with Rosannia. Lactoris fernandeziana of the monotypic family Lactoridaceae is endemic to Robinson Crusoe Island (formerly Masatierra Island) in the Juan Fernández Archipelago, Chile. © 2009 Elsevier B.V. All rights reserved.

1. Introduction Srivastava (1968a) instituted the genus Rosannia and described its type species Rosannia manika from the Edmonton Formation at East Coulee, Alberta (Fig. 1). Subsequently, Gibson (1977) raised the Edmonton Formation to group status and subdivided it into the Horseshoe Canyon, Whitemud, Battle, and Scollard Formations (Fig. 2). The Edmonton Group overlies the Bearpaw Formation and underlies the Paleocene Paskapoo Formation (Srivastava, 1994a). A number of radiometric dates have been published which help constrain the age of the formations. Lerbekmo and Braman (2002) published a date of 73.2 Ma for the Dorothy Bentonite, a unit that is in the upper part of the Bearpaw Formation. This suggests that the Horseshoe Canyon Formation should be about 73 my old (Fig. 2). Eberth and Deino (2005) provided a date of 70.44 Ma for an ash within the Drumheller Marine Tongue, a package sitting near the middle of the Horseshoe Canyon Formation. The Kneehills Tuff near the top of the Battle Formation has most recently been dated by Obradovich (1993) as 66.8 Ma. Using palynomorphs, the Cretaceous– Tertiary boundary has been placed at the base or near the base of a coal seam termed the Nevis coal that occurs near the middle of the Scollard Formation (Sweet et al., 1990; Sweet and Braman, 1991;

Srivastava, 1994a; Sweet and Braman, 2001). An ash at the top of the Nevis seam has been dated recently as 64.9 Ma (Eberth and Deino, 2005). The paleomagnetostratigraphy has been established for the entire section of the Edmonton Group from the Red Deer Valley to the Cypress Hills by Lerbekmo and Braman (2002). The Campanian– Maastrichtian boundary was placed near the base of the Drumheller Marine Tongue at the top of 32n magnetochron in Lerbekmo and Braman (2002) based on correlations with a section containing diagnostic ammonites in the Cypress Hills area of eastern Alberta. The radiometric date obtained by Eberth and Deino (2005) from the marine tongue is slightly above this placement of the boundary. Subsequently and independently, the stage boundary has been defined in Gradstein et al. (2004) at the top of 32n with a date of 70.6 Ma in the latest international time scale. The original diagnosis of the genus Rosannia and the description of its type species Rosannia manika were based on a misinterpretation of its morphological characteristics. Although the nomenclatural types of R. manika are well illustrated, the slide EC5/1 containing holotype and isotypes was loaned, never returned, and has presumably been lost. The purpose of this paper is to re-describe the type species, designate a lectotype and epitypes, and provide a revised diagnosis of the genus.

2. Material and method ⁎ Corresponding author. Tel.: +1 626 965 4014. E-mail addresses: [email protected] (S.K. Srivastava), [email protected] (D.R. Braman). 0034-6667/$ – see front matter © 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2009.10.003

All Rosannia manika type specimens were described (Srivastava, 1968a; Figs. 1–6 and 8) from sample EC5 of the East Coulee section

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Fig. 1. Map of southern Alberta showing sample collection localities for the Edmonton Group and the Dinosaur Provincial Park Formation.

except for an isotype (Srivastava, 1968a; Fig. 7) which is from sample DH39 of the Drumheller section. Both of these sections are from the lower part of the Horseshoe Canyon Formation (former lower Edmonton Formation) located along the Red Deer River in southern Alberta. The sampled interval is now thought to be latest Campanian in age. When a holotype is lost, a lectotype should be selected from the isotypes according to the International Code of Botanical Nomenclature (ICBN) (McNeill et al., 2006; Art. 9.10). Thus, the surviving isotype from sample DH39 of the Drumheller section (Srivastava, 1968a; Fig. 7) is here designated as a lectotype. Since the description of R. manika is being revised, epitypes are selected here to confirm the morphology of this pollen from slides EC5/II–IV of the same sample from which the holotype was originally designated. Five more translucent epitypes are illustrated to show various exine sculptures which are not otherwise apparent. Srivastava (1968b) measured and collected samples for palynological studies from the East Coulee and Drumheller sections. The East Coulee section is at the base of the Horseshoe Canyon Formation and in terms of coal seam nomenclature of Gibson (1977) would include coal seams 0 to 4. The Drumheller section includes coals 7 to 9 of Gibson (1977) and so there is an interval of non-overlap between the two sampled sections (Fig. 3). Srivastava's type material from the Edmonton Group is deposited in the permanent collections of the Royal Tyrrell Museum. All of the

specimens illustrated in this report are in these collections including the preparation of the recent Lactoris fernandeziana. Genus Rosannia S. K. Srivastava 1968a emend. S. K. Srivastava & Braman. 1968a — Rosannia S. K. Srivastava, p. 949. 1987 — Lactoripollenites Zavada & Benson, p. 1591. Type species: Rosannia manika S. K. Srivastava 1968a (monotypic)= Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava & Braman. Name derivation: Srivastava (1968a) coined the name of the genus Rosannia by combining Ros +ann +suffix ia after Rosalind Ann Catterall. Emended diagnosis: Pollen obligate tetrad, individual grains anaulcerate; ulcus large, round, ulcus edge smooth to uneven; exine calymmate, foot layer separates from baculate layer at the base of the pore forming a large pore chamber, called here cubiculum, infratectal ornamentation microreticulate; bacules infratectally arranged radially around the ulcus, supratectal ornamentation granulose. The term cubiculum is proposed here for a pore chamber formed by the separation of the baculate layer from the foot layer that forms the base of the chamber. An endopore may or may not be present. Thus, a cubiculum (L., pl. cubicula; meaning: sleeping room; bedchamber; cubicle) is defined as a chamber formed by abrupt separation of the foot layer and the baculate layer where the foot layer forms the base of the chamber. The terms atrium, fastigium or vestibulum describing

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Fig. 2. The age and classification of the Edmonton Group and its correlation with the Edmonton Formation. Column (1) gives the classification of the Edmonton Group (Gibson, 1977) and its overlying and underlying formations; (2) shows idealized composite lithological section of the Edmonton Group and its main lithological units (Srivastava, 1970); (3) shows ages of various units of the Edmonton Group (Lerbekmo and Braman, 2002; Eberth and Deino, 2005; Obradovich, 1993) and the range of Rosannia manika occurrence; (4) shows the classification of the Edmonton Formation (Allan and Sanderson, 1945) which was used in Srivastava (1968a).

various types of pore chambers (Punt et al., 2007) do not fully encompass the type of pore chamber dealt with here. Remarks: Srivastava (1968a) instituted the genus Rosannia but misdiagnosed it as a heteropolar, triprojectate, triporate pollen akin to heteropolar tricolpate genus Mancicorpus which occurs abundantly in the same assemblage. Its misdiagnosis was recognized for some time and it is revised here now. As revised the genus Rosannia includes obligate tetrads of ana-ulcerate pollen having supratectal granulose to scabrate ornamentation which is radially arranged around the pores. At the base of the cubiculum is a uniform membrane of foot layer in Rosannia which may or may not have an endopore. A “Saccus”-like thickening around the pore in Lactoripollenites (sensu Zavada and Benson, 1987) appears to be an artifact as a thin exine could sag, fold or invaginate and thus may give a thickened or “saccate” appearance at the pore with a large cubiculum. Zavada and

Benson (1987) considered the formation of the pore chamber in Lactoripollenites analogous to the saccus formation in gymnospermous bisaccate pollen by the sexine where the foot layer forms the base of the saccus. Macphail et al. (1999) pointed out that the saccus in gymnospermous pollen is filled with sexinous infratectal material whereas the “saccus” in Lactoripollenites is an empty chamber, i.e., devoid of any exinal material. Macphail et al. (1999) saw the need for a new morphological term for the pore chamber in Lactoripollenites. The term cubiculum may fill that need. Lactoripollenites includes obligate tetrads of monoaperturate anasulcate pollen with calymmate exine and an exinal thickening around the aperture forming a “saccus” (Zavada and Benson, 1987) which is here considered as a sagged exine at the pore. It has granulose supratectal sculpture. It cannot be distinguished from Rosannia and is considered here a junior synonym.

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The genus Walkeripollis Doyle, Hotton & Ward also has essentially similar characteristics but differs in having foveolate to finely reticulate tectal sculpture (Doyle et al., 1990). The genus Pseudowinterapollis Krutzsch differs in having large tectal reticulum as shown by its holotype (Krutzsch, 1970; holotype illustration: Couper, 1960, pl. 5, Fig. 4 in Raine et al., 2006). The genus Gephyrapollenites Stover & Partridge is a junior synonym of Pseudowinterapollis (Mildenhall and Crosbie, 1979). As shown by Zetter et al. (2002, pl. 8, Figs. 1, 2), Aachenipollis appears to be primarily ana-ulcerate but due to lateral compression the ulcus forms variable shapes like an elongated slit or a slit with rounded ends (pl. 8, Figs. 10, 11, 13, 14). The colpate aperture in the specimen of Aachenipollis aachenensis illustrated by Zavada and Benson (1987; Fig. 7) may be due to lateral compression of its large pore. Otherwise, Zavada and Benson's (1987) A. aachenensis specimen (Fig. 7) is very similar to their Lactoripollenites africanus paratype (Fig. 2). However, Aachenipollis is distinct from Rosannia in having acalymmate exine (Zetter et al., 2002). Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava & Braman. Plate I; Figs. 1–6; Plate II; Figs. 1-6. 1968a — Rosannia manika S. K. Srivastava, p. 949, Figs. 1–8. 1987 — Lactoripollenites africanus Zavada & Benson, p. 1591, Figs. 2–4. Holotype: Original designation (Srivastava, 1968a), size 32 μm; slide EC5/1, coordinates 45.7/116.3, Figs. 1, 2; East Coulee section, East Coulee (for sample position see Fig. 3), Horseshoe Canyon Formation, Alberta, Canada; slide is lost. Isotypes: Slide EC5/1, coordinates 61.0/122.8, Figs. 3, 4; EC5/1, coordinates 36.0/114.3, Figs. 5, 6; and EC 5/1, coordinates 65.4/113.0, Fig. 8 (Srivastava, 1968a); East Coulee section, Horseshoe Canyon Formation, Alberta, Canada; slide is lost. Lectotype: Plate I, 1, slide DH39/1, [coordinates 36.8/115.3, Srivastava (1968a)], Drumheller section, Drumheller (for sample position see Fig. 3); Horseshoe Canyon Formation, Drumheller, Alberta, Canada; an isotype designated in Srivastava (1968a, Fig. 7); England finder coordinates Q39/4; here designated as a lectotype and re-illustrated; specimen number TMP2008.222.0048. Epitypes: Plate I, 2, Slide EC5/IV, England finder coordinate G36/0, East Coulee section, Horseshoe Canyon Formation, East Coulee, Alberta, Canada, specimen number TMP2008.221.0022. Plate I, 3, slide EC5/II, England finder coordinate S48/3, East Coulee section, Horseshoe Canyon Formation, East Coulee, Alberta, Canada, specimen number TMP2008.221.0020. Plate I, 4, Slide EC5/III, England finder coordinate K28/0, East Coulee section, Horseshoe Canyon Formation, East Coulee, Alberta, Canada, specimen number TMP2008.221.0021. Plate I, 5, Slide Sc1-24/III, England finder coordinate D34/3, Scollard 1 section, Horseshoe Canyon Formation, Scollard, Alberta, Canada, specimen number TMP2008.226.0005. Plate I, 6, Slide 19b, sections 85–89, England finder coordinate W36/2, Dinosaur Park Formation, Dinosaur Provincial Park, Alberta, Canada, specimen number TMP1985.89.0063. Plate II, 1 and 2, Sample 13, Slide b, England finder coordinate M36/ 4, Battle River locality, Dinosaur Park Formation, Dinosaur Provincial Park, Alberta, Canada, specimen number TMP1989.270.0017. Plate II, 3 and 4, Sample 15, Slide b, England finder coordinate K63/ 1, Battle River locality, Dinosaur Park Formation, Dinosaur Provincial Park, Alberta, Canada, specimen number TMP1989.270.0018.

Fig. 3. East Coulee and truncated Drumheller sections. Srivastava (1968b, 1970) considered the top of the East Coulee section overlapping the base of the Drumheller section. Now it has been recognized that an interval of non-overlap from coal seam 4 to coal seam 6 (Gibson, 1977) exists between the two sections. The holotype of Rosannia manika occurred in Sample EC5 denoted by ⁎ and paratype [now designated lectotype] in sample DH39 denoted by ▲.

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Plate II, 5 and 6, Sample 74, Slide b, England finder coordinate L40/ 2, Willow Creek Section, Horseshoe Canyon Formation, Drumheller, Alberta, Canada, specimen number TMP1986.92.0030. Repository: Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada T0J 0Y0. Type locality: Drumheller outcrop section, Drumheller, Alberta, Canada. Stratigraphic horizon: Horseshoe Canyon Formation, Alberta, Canada; Campanian. Name derivation: Srivastava (1968a) derived the name from a Sanskrit word (adjective) manika meaning like a jewel. Mani is a mythical glowing jewel that only mature serpents could have as a source of light on their head to roam about in the dark to search for food. A person would be very lucky to possess it (nobody could get it even in mythology). The word is used for superlative qualities. Occurrence in Western Canada: Srivastava (1968a) reported Rosannia manika from the East Coulee and Drumheller sections of the Horseshoe Canyon Formation (Fig. 3). Subsequently it has been observed by Braman and Koppelhus (2005) throughout the Campanian Dinosaur Park Formation at Dinosaur Provincial Park, Alberta, that has been dated as 75 to 76.5 Ma (Eberth, 2005; Eberth and Deino, 2005). Lerbekmo et al. (2003) recovered it from the Bearpaw Formation in the Castor Well of central Alberta. Unpublished records of (DRB) show it is present in the Battle Formation and throughout the Horseshoe Canyon Formation along the Red Deer River in Alberta. It has not been found in the pre-Dinosaur Park Formation units in southern Alberta. Sweet and Braman (2001) reported the species from the Maastrichtian Summit Creek Formation at Police Island section in the Brackett Basin of the Northwest Territories of Canada. Previous records: Early Turonian to Campanian, southwest coast of western Africa (Zavada and Benson, 1987); Campanian–Eocene, Australia (Macphail et al., 1999); Senonian, Krishna–Godavari Basin, eastern India (Prasad et al., 1995); Santonian–Campanian, Cauvery Basin, eastern India (Venkatachala, 1974; Mehrotra et al., 2002, 2005; Prasad and Pundeer, 2002; Sastri et al., 1974; 1977; Venkatachala and Sharma, 1974); Campanian–early Maastrichtian, Alberta, Canada (Srivastava, 1968a; Braman and Koppelhus, 2005; Lerbekmo et al., 2003); Maastrichtian?–Danian and Early Miocene, Argentina (Ruiz and Quattrocchio, 1997; Quattrocchio and Ruiz, 1999; Quattrocchio et al., 2000; Gamerro and Barreda, 2008). Emended description: Pollen obligate tetrad, individual grains monoporate, ana-ulcerate, slightly constricted proximally but distally sometimes flared, folded or sagged around the pore; pore large, situated at the distal end of an individual grain (Plate I, 3); sexine calymmate (Plate I, 4; Plate II, 1, 4, 6), about 1.5 μm thick, tectate, tectum thin, baculate and foot layers of almost equal thickness, bacules short and wide, baculate and foot layers separate at the base of the pore forming a large cubiculum; endopore not seen; infratectal ornamentation microreticulate, formed by the arrangement of 4–5 bacules in a reticulate pattern (Plate II, 2, 5), lumina size less than 1.00 μm; bacules arranged radially on the pore area (Plate II, 5), arranged equatorially in the rest of the grain body; supratectal ornamentation granulose.

Measurements: Equatorial diameter 30–36 μm (Srivastava, 1968a); 34–36 μm (Zavada and Benson, 1987); 40–56 μm (Macphail et al., 1999); 30–32 μm (Gamerro and Barreda, 2008). Discussion and comparison: Rosannia manika includes obligate tetrads of monoporate ana-ulcerate pollen grains with calymmate exine. Pore or ulcus is large and circular with a diameter 20–23 μm (Plate I, 3). The cubiculum is large, occupying more than half of the individual grain (Plate I, 4). The cubiculum is formed by the separation of the foot layer from the baculate layer whereas the foot layer forms the base of the cubiculum and the baculate layer forms the rest of the cubiculum with a large opening as an ulcus (Plate I, 4). The SEM and TEM illustrations given by Zavada and Benson (1987; Figs. 2, 3) also show similar features in forming the cubiculum. The endopore occurrence is not clear. The TEM illustration of Zavada and Benson (1987; Fig. 3) also does not show a definite endopore. Exine of the ulcus area occasionally folds (Plate I, 1 and 6), or shreds at the end (Plate I, 5). A sagged or invaginated ulcus has been called a saccus by Zavada and Benson (1987). Zavada and Benson (1987) designated the holotype for Lactoripollenites africanus but did not illustrate it, although SEM, TEM and LM photomicrographs of the paratypes are illustrated (Figs. 2–4). Thus, for all practical purposes, their paratype replaces the holotype. As discussed above, the morphological characteristics of L. africanus are the same as those of Rosannia manika and hence the two species are considered herein synonyms. Botanical affinity: Lactoris fernandeziana Phil. of the family Lactoridaceae is known only to occur endemically on Robinson Crusoe Island (formerly Masatierra Island) in the Juan Fernández Archipelago, Chile (González and Rudall, 2001). The pollen morphology of extant L. fernandeziana has been compared in detail with Rosannia manika or with its synonym Lactoripollenites africanus (Zavada and Benson, 1987; Macphail et al., 1999; Gamerro and Barreda, 2008). Erdtman (1964, 1966) described the calymmate sexine and ulcus structure of L. fernandeziana. Due to the variable nature, its pore morphology remained elusive. With SEM, TEM, and OM studies, several morphological characteristics of extant Lactoris pollen can now be confirmed and compared more accurately with R. manika. The salient characteristics of R. manika and pollen of extant L. fernandeziana are compared below and are summarized on Table 1. Size: Srivastava (1968a), Zavada and Benson (1987), and Gamerro and Barreda (2008) reported the equatorial diameter of Rosannia manika tetrads ranging from 30 to 36 μm. The equatorial diameter of Australian specimens as documented by Macphail et al. (1999) is larger (40–56 μm). The equatorial diameter of extant Lactoris fernandeziana pollen also ranges from 30 to 36 μm (Erdtman, 1964, 1966; Sampson, 1995; Gamerro and Barreda, 2008) except that Heusser (1971) reported a size range from 50 to 70 μm. Exine: Rosannia manika has a calymmate exine, i.e., the sexine is continuous around the tetrad whereas the foot layer continues around each monad of the tetrad (Plate I, 1, 6; Srivastava, 1968a, Figs. 5, 6; Gamerro and Barreda, 2008, Figs. 11–14). Similar calymmate sexine occurs in Lactoris fernandeziana pollen (Plate III, 1–3; Erdtman, 1964, Fig. 1C; Gamerro and Barreda, 2008, Fig. 5). Calymmate sexine is very

Plate I. Bar scale = 10 μm. Fig. 1. Fig. 2. Fig. 3. Fig. 4. Fig. 5. Fig. 6.

Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, lectotype, TMP2008.222.0048, DH39/I, Q39/4, Drumheller Section, Drumheller, Alberta, Horseshoe Canyon Formation; showing infratectal microreticulate sculpture. Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, EC5/IV, TMP2008.221.0022, G36/0, East Coulee Section, East Coulee, Alberta, Horseshoe Canyon Formation; showing infratectal microreticulate sculpture and calymmate exine (indicated by an arrow). Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, EC5/II, TMP2008.221.0020, S48/3, East Coulee Section, East Coulee, Alberta, Horseshoe Canyon Formation; shows folded exine at the distal end of a monad. Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, EC5/III, TMP2008.221.0021, K28/0, East Coulee Section, East Coulee, Alberta, Horseshoe Canyon Formation; showing an ulcus in optical view with folded exine. Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, Sc1-24/III, TMP2008.226.0005, D34/3, Scollard locality 1, Scollard, Alberta, Horseshoe Canyon Formation; showing well developed folds on ulcus margins. Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, TMP1985.89.0063, Sample 19, slide b, W36/2, Dinosaur Provincial Park, Alberta, Dinosaur Park Formation; showing petaloid exinal folds around the ulcus.

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distinctly seen in a TEM section of an extant Lactoris pollen illustrated by Sampson (1995, Fig. 2A). Sexine is baculate with short and wide bacules in Rosannia manika (Plate I, 2–5; Plate II, 1, 2, 5, 6). The short and wide nature of bacules is well illustrated in a TEM section of a specimen from southern Africa (Zavada and Benson, 1987, Fig. 3). A few simplicolumellate strands run from the base of the cubiculum to its distal end (Plate I, 3; Plate II, 1). Similar sexine is shown in the TEM sections of extant Lactoris pollen (Zavada and Benson, 1987, Fig. 6; Sampson, 1995, Fig. 2F). Tectal ornamentation is microreticulate in Rosannia manika. Four or five bacules join each other with a lumina in between them (Plate I, 1 (lectotype); Plate II, 2, 5; Srivastava, 1968a, Figs. 1 (holotype), 4, 6, 8). Heusser's (1971) foveolate exine and Zavada and Benson's (1987, Fig. 3) “tectal perforations” in between bacules in Lactoris pollen are homologous to microreticulate sexine of R. manika (Srivastava, 1968a). Similar minute perforations are demonstrated in the TEM sections of extant Lactoris pollen (Zavada and Benson, 1987, Fig. 6; Sampson, 1995, Fig. 2F). Supratectal ornamentation is scabrate in Rosannia manika (Plate I, 1, lectotype; Srivastava, 1968a, Fig. 1, holotype; the SEM illustration — Zavada and Benson, 1987, Fig. 2). Similar scabrate ornamentation occurs in extant Lactoris pollen as seen in published SEM illustrations (Walker, 1976, Fig. 9:4; Zavada and Benson, 1987, Fig. 5; Sampson, 1995, Figs. 1B–I). Some Rosannia manika specimens have exinal folds proximally on the monads of the tetrad (Srivastava, 1968a, Figs. 3, 5, isotypes; Zavada and Benson, 1987, Fig. 2, SEM illustration). Similar folds occur in extant Lactoris pollen also (Walker, 1976, Fig. 9:4; Sampson, 1995, Fig. 1F). Pore structure: Rosannia manika as well as extant Lactoris fernandeziana pollen have ana-ulcerate monoporate monads of obligate tetrads. R. manika pores are large and circular (Plate I, 2, 3, 6; Srivastava, 1968a, Figs. 5, 6, isotype). The pore diameter in some specimens extends across the whole distal end of the monad (Plate I, 1, lectotype; Srivastava, 1968a, Fig. 1, holotype). The foot layer separates from the sexine at the base of the pore and forms the base of a large cubiculum (Plate I, 2). A definite endopore has not been seen in R. manika. Distally the cubiculum may be flared, folded or sagged (Plate I, 1, 5, 6; Zavada and Benson, 1987, Fig. 2). Pore morphology in extant Lactoris pollen is variable and controversial. Zavada and Taylor (1986) and Zavada and Benson (1987) considered the sagged cubiculum as a “saccus” and compared the separation of the foot layer from the sexine at the base of the cubiculum with that of the saccus in gymnosperm pollen. The formation of a cubiculum in Lactoris pollen is similar to the formation of any other pore chamber in angiosperm pollen by the separation of foot layer and sexine, such as an atrium or fastigium. Erdtman (1964, Fig. 1B) illustrated a specimen of Lactoris pollen having a circular pore with a diameter as large as the width of the monad which compares well with Rosannia manika (Plate I, 3). Heusser (1971, p. 39) described the pore area as “… distally wrinkled and flanged appearing somewhat knob-like with a recessed, more or less circular, pore-like area or tenuitas, ca 17 μm across”. Flared, folded or sagged distal cubiculum is also seen in some extant Lactoris pollen (Plate II, 6; Zavada and Taylor, 1986, Figs. 1, 5, SEM illustrations; Zavada and

Benson, 1987, Fig. 5, SEM illustration). Sampson (1995) concluded that Lactoris pollen from preserved flowers had a small poorly defined aperture (Fig. 1B, SEM illustration) but had a collapsed aperture after the emergence of the pollen tube (Fig. 1I, SEM). However, we found both types of pollen occurring in a slide of the herbarium material of Lactoris pollen (Plate III, 2–6). The comparison of the exine, pore structure, infratectal and supratectal sexine ornamentation indicates a close affinity between Rosannia manika and extant Lactoris fernandeziana pollen. The salient characteristics of R. manika and extant L. fernandeziana pollen are summarized on Table 1 for comparison. Phytogeography of Rosannia manika: The extant Lactoridaceae is a monotypic family represented by Lactoris fernandeziana which is endemic to Robinson Crusoe Island (formerly Masatierra Island). This island, 667 km west of the Chilean coast of South America in the Pacific Ocean, forms the eastern part of the Juan Fernández Archipelago. The western part of the archipelago is formed by Alejandro Selkirk Island (Musafuera Island). Also within the archipelago is Santa Clara Island near Robinson Crusoe Island. The Archipelago is volcanic in origin (Greimler et al., 2002). The radiometric ages of Robinson Crusoe Island and Alejandro Selkirk Island are about 4 my and 1–2 my, respectively (Stuessy et al., 1984). Robinson Crusoe Island is mountainous with rugged coastlines and a subtropical climate. L. fernandeziana is a shrub, about 1 m tall, with brittle stems and obovate leaves [broader at the apex and narrower at the base] (Stuessy et al., 1998). It is restricted to “windy fog- and rain-swept mountain forests on steep slopes” of Robinson Crusoe Island (Bernardello et al., 1999). Fossil records of Lactoridaceae indicate that it originated in Turonian sediments, about 92–90 my BP, in the Orange Basin offshore Namaqualand, South Africa (Zavada and Benson, 1987). The type material was documented from cutting samples. Such samples are notorious for reworking from younger horizons so only top occurrences are used reliably in correlations and age determinations. Thus, the Turonian age for Lactoridaceae should be accepted cautiously until more definite proof comes forward for its antiquity. Still, the migration of Lactoridaceae from South Africa to its present abode in an offshore island west of Chile is intriguing. Fossil Lactoridaceae occurs in Australia from Campanian to early Oligocene (Macphail et al., 1999). The lack of geological evidence for any land connection between Africa and Australia during the Late Cretaceous (Metcalfe, 2001) presents difficulty in explaining how the Lactoridaceae reached Australia from Africa. Macphail et al. (1999) considered that the Lactoridaceae could have entered Australia only through northern Australia from Africa during the middle-late Campanian, flourished on the continent through the hot climate during the latest Paleocene–early Eocene, and became extinct there in the Oligocene of Zeehan district, western Tasmania, with the approach of a cold climate indicated by Oligocene tillites. Lactoridaceae is present in the Senonian of India (Prasad et al., 1995; Prasad and Pundeer, 2002) and South America (Ruiz and Quattrocchio, 1997). Palynological and paleontological data indicate that India was connected with Africa in the Senonian (Srivastava, 1988; Briggs, 2003). Africa and South America also had land connections across the Brazilian bulge with Nigeria until the end of

Plate II. Scale bar = 10 μm. Figs. 1 and 2.

Figs. 3 and 4. Figs. 5 and 6.

Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, TMP1989.270.0017, Sample 13, slide b, M36/4, Battle River, Dinosaur Park Formation. Figures show different levels of focus. Arrow in Fig. 1 indicates calymmate exine. Arrow in Fig. 2 is directed to area where reticulations are visible. Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, TMP1989.270.0018, Sample 15, slide b, K63/1, Battle River, Dinosaur Park Formation. Figures show different levels of focus. Arrow in Fig. 4 indicates calymmate exine. Rosannia manika S. K. Srivastava 1968a emend. S. K. Srivastava and Braman, TMP1986.92.0030, Sample 74, slide b, L40/2, Willow Creek Section, Horseshoe Canyon Formation, Drumheller. Figures show different levels of focus. Arrow in Fig. 5 is directed to area where reticulations are visible. Bacules on the margin show narrow stems and broader heads (capita). Arrow in Fig. 6 indicates calymmate exine.

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Table 1 Comparison summary of the salient characteristics of Rosannia manika and extant Lactoris fernandeziana pollen. Characteristics

Rosannia manika

Pollen of extant Lactoris fernandeziana

Size: diameter of tetrads

30–36 mm (Srivastava, 1968a; Zavada and Benson, 1987; Gamerro and Barreda, 2008); 40–56 mm in Australian specimens (Macphail et al., 1999). Sexine calymmate, foot layer appears to mark the boundaries of monads of the tetrads (at the lower angular point; Plate I, 4, 6) but not clear (Srivastava, 1968a, Figs. 5, 6; Gamerro and Barreda, 2008, Figs. 11–14). Sexine baculate, bacules short and wide. A few strands run from the base of the cubiculum to its distal end, strands simplicolumellate (Srivastava, 1968a, Figs. 1–8; Plate I, 2–5; Plate II, 1, 2, 4, 5; Zavada and Benson, 1987, Fig. 3). Reticulate, lumina less than 1.00 mm (Srivastava, 1968a; Plate I, 1–3; Plate II, 1–6; Zavada and Benson, 1987, Fig. 3, TEM; Gamerro and Barreda, 2008, Figs. 13, 14). Granular or scabrate (Plate I, 1, 4; Plate III, 1, 2, 5; Srivastava, 1968a, Figs. 1, 5, 7; Zavada and Benson, 1987, Fig. 3, TEM). Ana-ulcerate, large, situated at the distal end of the cubiculum formed by the separation of sexine and foot layer where foot layer forms the base of the cubiculum (Plate I, 4; Zavada and Benson, 1987, Fig. 3, TEM; Gamerro and Barreda, 2008, Figs. 11–15). Ulcus small and circular (not reported).

30–36 mm (Erdtman, 1964, 1966; Sampson, 1995; Gamerro and Barreda, 2008); 50–70 mm (Heusser, 1971).

Exine

Sexine sculpture

Tectal ornamentation Supratectal ornamentation Ulcus morphology

Sagged or folded ulcus (Plate I, 1, 4–6; Plate II, 3–4; Zavada and Benson, 1987, Fig. 2, SEM; Macphail et al., 1999, Figs. 2–38).

the Cretaceous (Ojeda, 1982; Rand and Mabesoone, 1982). The Late Cretaceous Constantinisporis phytogeoprovince (Srivastava, 1994b) across Africa, South America and India had several pollen genera common in the three continents. Thus, Lactoridaceae could have migrated west from Africa to South America (Ruiz and Quattrocchio, 1997) and east to India (Prasad et al., 1995; Prasad and Pundeer, 2002) during the Senonian. The insertion of the Caribbean Plate from the Pacific in the Campanian (Pitman et al., 1993) connected North America with South America and opened up a land-plant migration path between the two continents in the Senonian. The Lactoridaceae flourished during the Campanian and early Maastrichtian in southern Alberta, Canada (Fig. 2). The deltaic Horseshoe Canyon Formation, deposited on the western coast of the North American inland sea, supported large Taxodium forests. The prevailing climate was humid subtropical, similar to the present-day climate in the southeastern coastal plain in North America at latitude 25–30°N (Srivastava, 1994b). The humid subtropical climate started changing to a warm temperate one during the deposition of the Whitemud Formation (Fig. 2) at the top of the Horseshoe Canyon Formation. The last occurrence of Lactoridaceae in North America is in the early Maastrichtian in the Battle Formation near the top of the Horseshoe Canyon Formation of Alberta. The youngest fossil record of Lactoridaceae is in the early Miocene Austral Basin, Argentina, which was deposited under a prevailing temperate to warm temperate climate (Gamerro and Barreda, 2008). This site is about 2000 km across the high Andes mountains southeast of

Sexine calymmate, foot layer marks the boundaries of tetrad monads (Erdtman, 1964, Fig. 1C; Sampson, 1995, Fig. 2A, TEM section; Gamerro and Barreda, 2008, Fig. 5). Sexine baculate, bacules short and wide (Zavada and Benson, 1987, Fig. 6; Sampson, 1995, Fig. 2F).

Tectum having small perforations (reticulation) (Plate III, 5; Heusser, 1971; Zavada and Benson, 1987, Fig. 6, TEM; Sampson, 1995, Fig. 2F, TEM). Granular (Zavada and Benson, 1987, Fig. 6, TEM; Sampson, 1995, Figs. 1B–1I). Ana-ulcerate, large, situated at the distal end of the ulcus chamber (cubiculum) formed by the separation of sexine and foot layer where foot layer forms the base of the cubiculum (Zavada and Benson, 1987, Fig. 6, TEM; Gamerro and Barreda, 2008, Figs. 3–7). Ulcus small and circular (Plate 3, Fig. 21; Erdtman, 1964, Fig. 1A; Sampson, 1995, Fig. 1B). Sagged or folded ulcus (Plate III, 17; Heusser, 1971, Fig. 32–390; Zavada and Benson, 1987, Fig. 5, SEM; Sampson, 1995, Fig. 1I; Macphail et al., 1999, Figs. 44–48; Gamerro and Barreda, 2008, Figs. 8–10).

the present abode of Lactoridaceae on Robinson Crusoe Island, west of the Chilean coast. Gamerro and Barreda (2008) considered that Lactoris migrated to the west before the Andean maximum uplift in the middlelate Miocene. The fossil record of Lactoridaceae is far from complete in South America making it impossible to map its migration path accurately. Considering the Senonian land connection between Nigeria and Brazil (Ojeda, 1982; Rand and Mabesoone, 1982), Lactoridaceae could have entered southern South America from the north. Lactoridaceae appears to have adapted now to cooler and harsher climatic conditions than during its heyday in the warm Late Cretaceous. Its arrival on Robinson Crusoe Island is considered rather recent as the island is only about 4 my old (Stuessy et al., 1984). Gunnera is another genus associated with the Lactoris plant community on this island (Pacheco et al., 1991; Greimler et al., 2002). The paleophytogeography and migration pattern of Gunnera (Srivastava, 1994b; Wanntorp et al., 2004) are very similar to those of Lactoris. Late Quaternary profiles from Alexander Selkirk Island show that Gunnera reached there at least 14,000–12,000 14C yr BP and is extant on that island (Haberle, 2003). Lactoris pollen was not found in these samples. A similar study has not been done for the older Robinson Crusoe Island. Gamerro and Barreda (2008) hypothesized several ways that Lactoris could have crossed the water barrier between the Chilean mainland and Robinson Crusoe Island. The low sea level during the glacial interval about 2 my BP may have opened up a land connection between the mainland and Robinson Crusoe Island (Gamerro and Barreda, 2008) facilitating the migration of Lactoris.

Plate III. Bar scale = 10 μm. Fig. 1. Fig. 2. Fig. 3. Fig. 4. Fig. 5. Fig. 6.

Lactoris fernandeziana Phil., Robinson Crusoe Island (formerly Masatierra Island), Juan Fernández Archipelago, Chile, TMP2008.219.0001, P38/0; a tetrad. Lactoris fernandeziana Phil., Robinson Crusoe Island (formerly Masatierra Island), Juan Fernández Archipelago, Chile, TMP2008.219.0002, M36/4; a tetrad showing pore distally. Lactoris fernandeziana Phil., Robinson Crusoe Island (formerly Masatierra Island), Juan Fernández Archipelago, Chile, TMP2008.219.0003, P36/3; another tetrad showing an ulcus at the distal end of the top monad. Lactoris fernandeziana Phil., Robinson Crusoe Island (formerly Masatierra Island), Juan Fernández Archipelago, Chile, TMP2008.219.0004, K38/4; showing a well formed pore at the distal end of the top monad. Lactoris fernandeziana Phil., Robinson Crusoe Island (formerly Masatierra Island), Juan Fernández Archipelago, Chile, TMP2008.219.0005, O38/1; showing a pore in the top monad. Lactoris fernandeziana Phil., Robinson Crusoe Island (formerly Masatierra Island), Juan Fernández Archipelago, Chile, TMP2008.219.0006, N38/0; a tetrad showing folded exine at the margin of the ulcus.

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Acknowledgements Lucy M. Cranwell was the first to draw SKS' attention to the similarity between Rosannia manika and extant Lactoris pollen soon

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after the publication of Srivastava (1968a). She repeatedly urged him to publish on the affinity of Rosannia but regrettably it was delayed until now. Viviana Barreda provided a pollen slide of the herbarium material of Lactoris fernandeziana Phil. which is used here for

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